54 | : Beet EMITASON/ AT ENT MAY 102004 } Volume 58 Number 1 22 April 2004 N\wer aries ISSN 0024-0966

Journal of the

Lepidopterists Society

Published quarterly by The Lepidopterists’ Society

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Cover illustration: Two late-instar Monarch (Danaus plexippus (L.): Nymphalidae) caterpillars “face off’ over the last bit of Milkweed (Asclepias) leaf, Northern Wisconsin, USA. Photo by Michael Toliver.

JOURNAL OF

Tue LeEpipopTeERISTS’ SOCIETY

Volume 58

Journal of the Lepidopterists’ Society 58(1), 2004, 1-6

2004 Number |

SEASONAL DYNAMICS OF LEAF-TYING CATERPILLARS ON WHITE OAK

JOHN T. Litt! Department of Biology, University of Missouri-St. Louis, 8001 Natural Bridge Road, St. Louis, Missouri 63121, USA

ABSTRACT. This study examined the seasonal pattern of leaf tie construction and occupancy by caterpillars associated with white oak, Quercus alba, in Missouri. Thirty white oak trees were monitored over the course of a season, recording the phenology of leaf tie construction, occupancy, and abandonment by 14 different species of leaf-tying caterpillars. Larvae of Pseudotelphusa sp. (Gelechiidae) created most of the initial leaf ties, many of which were secondarily occupied by a variety of other leaf tiers. In addition, non-tying caterpillar species from a variety of families were common secondary occupants. As many as 6 different species and 15 different individual larvae were found occupying a single leaf tie over the course of the season. In selecting pre-existing leaf ties for colonization sites, the leaf tiers did not discriminate between ties of different ages. On average, one-third of the leaves on a branch were incorporated into a leaf tie at some time during the season. Despite their

abundance, leaf ties in this system appear to be a limiting resource.

Additional key words:

Shelter-building caterpillars are a conspicuous com- ponent of the insect herbivore fauna attacking temper- ate broad-leaved trees (Frost 1959, Prentice 1965). A variety of caterpillars in at least 12 (Berenbaum 1999) and as many as 27 (Jones 1999) different families use silk to construct leaf ties, webs, folds, and rolls that typically serve as both shelters and feeding sites (room and board) for developing larvae. These leaf shelters are often colonized subsequently by other arthropods, including other shelter-building caterpillars (Carroll & Kearby 1978, Cappuccino 1993, Cappuccino & Martin 1994, Lill 1999, Martinsen et al. 2000, Lill & Marquis 2003), many of whom oviposit in existing shelters (Lill & Marquis 2004). In the case of leaf ties (“sandwiches” of overlapping leaves), there is often a high rate of sec- ondary occupation by both leaf-tying and non-leaf-ty- ing caterpillars (Carroll et al. 1979, Fukui 2001) result- ing in multiple individuals of the same or different species co-occuring within a leaf tie. Such secondary occupation suggests that these shelters represent a po- tentially limiting resource for populations of leaf tiers and leaf tie associates and may, in part, influence the local abundance of caterpillars occupying individual host plants.

‘Current Address: George Washington University, Department of

Biological Sciences, 340 Lisner Hall, 2023 G Street, NW, Washing- ton, DC 20052, USA, email: lillj@gwu.edu

Leaf tie, insect phenology, positive interactions, Quercus alba, shelter-builders.

An important first step in understanding the dy- namics of these interactions is to document the sea- sonal pattern of shelter construction (i.e., resource availability), maintenance, and occupancy (resource use) by different species of caterpillars. Here, I de- scribe the natural history of leaf tie construction and occupancy of the leaf ties found on saplings of white oak (Quercus alba L.).

MATERIALS AND METHODS

This study was conducted in east-central Missouri at Cuivre River State Park. The park consists of second growth oak-hickory forest dominated by a canopy of white oak (Q. Glyn. black oak (Q. velutina Lam.), and hickory (Carya spp.) with an understory of flowering dogwood (Cornus florida L.), sassafras (Sassafras al- bidum Nees), redbud (Cercis canadensis L.), sugar maple (Acer saccharum Marsh) and various oak saplings. The study was conducted within the Big Sugar Creek watershed, an unmanaged natural area of the park.

Thirty small white oak trees (understory saplings) were tagged in early spring of 1996, prior to budbreak. Trees raced in height from 2 to 4 m and had at least four accessible branches randomly assigned to one of two treatments, control and census (two branches each per tree). Leaf ties formed on census branches were opened regularly to record the occupants whereas leaf ties on control branches were left undisturbed. The to-

bo

aK JURNAL OF THE LEPIDOPTERISTS’ SOCIETY

12 10 oO o G 8 Oo —®— New ties oS 6 —O— Abandoned ties > —v— Net ties 2 —v— Leaf-tying caterpillars =}. o) S Fn 2 2 ® Q 0

t So

5/27 6/10 6/24 7/8 7/22 8/5 Census date

Fic. 1.

tal number of leaves on each branch was recorded fol- lowing budbreak. On 29 May, I established three per- manent artificial leaf ties on one control and one census branch of each tree by clipping together haphazardly selected clusters of three adjacent leaves with light- weight curler clips (one clip/three-leaf cluster; Brent- wood Beauty Labs International, Hillside, Illinois). Ar- tificial leaf ties were created to sample the community of secondary occupants in a standardized manner (all artificial ties were initially unoccupied and established on the same date). Beginning 29 May, the number of all naturally occurring leaf ties present on each branch during eight census weit (29 May, 11] June, 20 June, 9 July, 23 July, 12 August, 27 August, and 14 Septem- ber) was recorded. In addition, for all branches in the census treatment, the contents of the leaf ties (both natural and artificial) were recorded by briefly opening the ties and then clipping them back together with a curler clip. Any caterpillars that escaped ion the ties by spinning down on silk were placed back into the leaf tie. The number and species of all leaf-tying cater- pillars and non-tying associates (e.g., herbivorous bee- tles and non-tying caterpillars) were recorded during each census. Each leaf tie in the census treatment was assigned a unique label so the sequence of occupancy (er individual ties) could be followed. Clips were re- moved at subsequent censuses from any natural leaf ties that were not actively maintained (i.e., that came apart upon removal of the clip or that contained no leaf-tying caterpillars during two consecutive cen- suses). None of the leaf ties on control branches were opened, but the artificial ties on these branches were externally inspected for signs of occupancy (e.g., skele- tonization damage, visible silk strands) to determine the timing of colonization.

Ss LSSLSS nA aS |

8/19 9/2 9/16

Seasonal pattern of natural leaf tie formation and leaf-tying caterpillar density on white oak. Error bars are 1 SE.

The minimum number of total individuals and species of leaf-chewing insects (caterpillars, beetles, and sawfly larvae) occupying a leaf tie over its “life- span” was determined by examining the sequence of occupation of each natural leaf tie; consecutive records of the same species were always assumed to be a single individual, so these estimates were likely to be conser- vative. The relationship between the age of the leaf tie (expressed as the number of censuses in which it was maintained) and the total minimum number of species and individuals was examined with linear regression. In addition, for leaf ties maintained for a given number of censuses (3, 4, or 5), the effect of the date of tie origination on the total minimum number of species and individuals was examined graphically.

Because the attractiveness of leaf ties to potential colonists might be expected to change as a tie ages (ac- cumulating damage, frass, and silk), I examined the in- fluence of tie age on the likelihood of secondary colo- nization by leaf-tying caterpillars. The distribution of “new” colonization events (occupancy by one or more individuals of a species not present in the previous cen- sus) across natural ties of different ages was compared with a null distribution using a Chi-square test. The null distribution assumed that ties of different ages would be colonized in proportion to their relative availability (i.e., colonists would not discriminate among ties of differ- ent ages). Because late season colonists had the widest range of tie ages available to “choose” from, only the new colonization events that occurred during each of the last two censuses were considered in this analysis.

RESULTS

Leaf ties. The total abundance of naturally occurring leaf ties (census + control treatments) ranged from a

VOLUME 58, NUMBER 1

TaBLE 1. Leaf-tying caterpillars colonizing white oak trees.

Peak density

Family and species* (larvae/100 Ivs.)

Gelechiidae Arogalea cristifasciella Chambers 0.30 Chionodes fuscomaculella Chambers 0.30 Pseudotelphusa sp. (undescribed) 7.45 Coleotechnites quercivorella Chambers 0.01 Noctuidae Morrisonia confusa Hubner 0.06 Oecophoridae Antaeotricha humilis Zeller 0.03 Antaeotricha osseella Williamson 0.01 Antaeotricha schlaegeri Zeller 0.17 Psilocorsis quercicella Chambers 1.86 Psilocorsis cryptolechiella Chambers 2.64" Psilocorsis reflexella Clemens — Setiostoma xanthobasis Zeller 0.04 Pyralidae Tetralopha expandens Walker 0.23 Tortricidae Anclis divisana Walker 0.07

*Nomenclature and authors follow Hodges (1983). Density data for P. cryptolechiella and P. reflexella are combined because early instars of these two species could not be distinguished.

low of 154 on 29 May (census 1) to a high of 1265 on 9 July (census 4). However, because branches varied in size (mean + 1 SE = 107.6 + 6.0 leaves), tie densities (no. leaf ties/leaf) were used in all analyses. Census branches and control branches did not differ from each other in the mean density of leaf ties in any of the eight censuses (p > 0.10 for all paired t-tests), suggesting that the disturbance and use of clips in the census treatment did not adversely affect tie formation patterns.

The rate of leaf tie formation (mean number of new leaf ties per census branch) increased markedly be- tween mid- and late June and peaked in early July dur- ing which time an average of four new leaf ties were formed per 100 leaves on each of the 60 census branches (Fig. 1). Following this peak, the rate of tie formation declined markedly while the number of abandoned ties (those that were empty and/or not maintained) increased, resulting in a decline in the net number of leaf ties present on the branches. During the last two censuses (late August and mid-September), the rate of new tie formation increased slightly while the number of abandoned old ties declined rapidly, re- sulting in a second peak in net tie density.

The total density (accumulated over the season) of natural leaf ties formed on a branch in the census treatment varied from 6.4 to 34.6 ties/100 leaves (mean + 1 SE = 17.6 + 0.80). Since most leaf ties con- sisted of two leaves, this means that on average, slightly more than one-third (17.6 + 2 = 35.2/100) of the leaves on a given branch were incorporated into a leaf tie at some point during the season. Because nat-

(ew)

ural leaf ties on the control branches were not fol- lowed individually, their seasonal pattern of tie forma- tion, occupancy, and abandonment could not be exam- ined. The total density of natural leaf ties formed on a census branch over the course of the study was posi- tively related to the density of initial leaf ties formed early in the season (i.e., by the second census on June 11; r? = 0.16, F, = 10.89, p = 0.002).

Occupants. Over the course of the season, 14 dif- ferent species of leaf-tying caterpillars (Table 1) were recorded as occupants of the naturally occurring leaf ties (N = 1162) on the census branches. Across the season, the density of leaf-tying caterpillars (all 14 species combined) varied from a low of 1.49 to a high of 9.50 larvae/100 leaves (Fig. 1). The density of leaf tiers peaked in late June and again in late August, slightly in advance of the “cells in leaf tie density. Among the seven most common species of leaf tiers, Pseudotelphusa sp. (Gelechiidae, currently unnamed) achieved the highest peak density (Table 1), and its two population peaks (representing its two genera- tions) preceded the peaks of the other bivoltine species (Fig. 2). Three species of Oecophoridae in the genus Psilocorsis (P. cryptolechiella, P. quercicella, and P. reflexella) were the next most prevalent, with peak densities >1 per 100 leaves.

Individual leaf ties varied considerably in the num- ber of species of leaf tiers colonizing them, ranging from a low of zero (for ties that had been created and abandoned prior to a census) to as many as six differ- ent species occurring in a single tie over its lifespan. The total number of individuals occupying a leaf tie over its lifespan varied from 0-15 (mean + 1 SE = 2.33 + 0.06). Within a particular census, it was not uncom- mon to find assemblages of several individuals from 2-3 different species sharing a leaf tie (the record was 9 caterpillars of 3 different species).

Leaf ties that hosted both a greater abundance and higher species richness of caterpillars were maintained for longer periods (Fig. 3). However, most ties were relatively ephemeral, lasting on average 3.3 + 0.1 cen- suses (a little more than a month). Of 91 natural ties recorded on census branches during the first census, only 10 (11%) persisted until the last census. However, for those natural ties (N = 302) formed during the first peak of leaf tie construction (9 July, census 4), almost a third (27%) were still occupied on the last census (14 September), which was near the second peak in leaf tie construction (see Fig. 1). The date that a leaf tie was formed also influenced the total abundance and species richness of colonists. For leaf ties that per- sisted for an equal number of censuses (either 3, 4, or

5), the average abundance and species richness tended

4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY = n 0 > w ® 8 = — ——@—— Pseudotelphusa sp. ey Oc). Lee / Pay, SS By le UA ° Psilocorsis reflexella c KA e) / ? yp ACA / ———¥ —— Psilocorsis quercicella = Zip, fares + xs ee — -7-—-+- Arogalea cristifasciella = ae a ra Ses ag a / —-#— _ Tetralopha expandens 2 0.01 4 He / j Pa / / ——O--— _— Chionodes fuscomaculella oO nee =e 5 5 if fo Antaeotricha schlaegeri e 0 oll / \ aut 2 coin] ge ney eines ms 0014 2 9 i | 7 \ ; / 3) sas heat / Hf / / : / 0.0001 ++ hia a an rm f AEA Ae 5/27 6/10 6/24 7/8 7/22 8/5 8/19 9/2 9/16 Census date Fic. 2. Seasonal dynamics of seven species of leaf-tying caterpillars on white oak. New ties are ties that had not been recorded in a previous

census; abandoned ties were not maintined and came apart readily upon examination; net ties = ties present in the previous census + new ties

— abandoned ties. Note log scale of ordinate.

to increase with later date of origination (Fig. 4), sug- gesting that new ties formed later in the season are subject to higher levels of secondary colonization than those formed earlier.

As the season progressed, heavily utilized leaf ties became highly damaged and accumulated frass from previous occupants, decreasing the amount of food available for developing larvae. Despite this apparent decline in resource quality, the age of the leaf tie had no influence on the probability of colonization by leaf tying caterpillars late in the season. For old ties (ties that had been established at least one census prior), the probability of a new colonization event was unre- lated to tie age (i.e., the observed distribution of colo-

7 6 4 54 o = g 4 € 3 iS w 3] fe) = 2 4 14 s @ Species © Individuals o+ Tos T ares US Samra ina T aaa 1 2 3 4 8 6 7 8

Age of leaf tie (no. of censuses)

Fic. 3. Total number of individuals and species of leaf-chewing insects recorded in leaf ties maintained for variable lengths of time. Each data point is a mean of a different set of leaf ties (data are not cumulative).

nization events across ties of different ages was not dif- ferent from the null expectation in either of the last 2 censuses; census 7: X°, , = 3.49, p > 0.50, N = 299; cen- sus 8: x7). ='9.17, p> 0.10; N = 119):

An additional 342 non-tying individuals, including 12 species of Lepidoptera, two leaf-feeding beetles, and one sawfly larva, were also found inhabiting the ties (Table 2). Non-herbivorous inhabitants were not recorded, but included click beetles, psocids, rove beetle adults and larvae, shield bugs, thrips, and lacewing larvae. Spiders often occupied the ties as well, occasionally making nests (with egg masses) in- side the ties.

Artificial ties. Artificial ties were readily colonized both by leaf-tying caterpillars and by non-tier asso- ciates. By late June, ninety-five percent of the 180 artificial leaf ties had been colonized by leaf tiers and all remaining ties were colonized by late July. For the 90 artificial ties on the census branches, a total of 733 leaf tier caterpillars and 202 non-tier associates were recorded during the seven censuses. All leaf tier species listed in Table 1 except Anclis divisana and Tetralopha expandens colonized the artificial ties. The density of leaf-tying caterpillars occupying these artificial leaf ties was similar to that observed for the natural ties.

DISCUSSION

There was marked seasonal variation in the con- struction of new leaf ties. Because Pseudotelphusa sp. larvae were the first leaf tiers to appear, they were largely responsible for the initial wave of ties, which were subsequently colonized by the later- appearing species (Fig. 2). The increased densities

VOLUME 58, NUMBER 1

3.5 Duration = 3 censuses

|

254

205

Total number/tie

5/27 6/10 6/24 7/8 7/22 8/5

Duration = 4 censuses

Total number/tie nN oO 4 KoH Hey OH He

0.5 5/27 6/10 6/24 718 7/22 8/5

se AL ee

5.0 = ] Duration = 5 censuses 454

Total number/tie w i=)

15 4 e@ Species © Individuals

5/27 6/10 6/24 7/18 7/22 8/5

Tie origination date

Fic. 4. Total number of individuals and species of leaf-chewing in- sects recorded in leaf ties originating on different dates, but main- tained for equal periods of time (3-5 censuses). Error bars are 1 SE.

of leaf tiers relative to leaf ties during the early sea- son (Fig. 1) could be explained by the fact that most of these early leaf ties were created by small groups (2-3) of Pseudotelphusa sp. larvae. The lack of a sec- ond peak in the production of new leaf ties during the late season (i.e., when leaf-tying caterpillar abun- dance reached its second peak) indicated that many of the new colonists (including the second wave of

UL

TaBLE 2. Non-leaf-tying herbivores (leaf-chewers) found inside leaf ties. Order Family Species* Coleoptera Chrysomelidae — Pachybrachis sp. Curculionidae — Cyrtepistomes castaneus Roelfs Hymenoptera Tenthredinidae — Caliroa sp. Lepidoptera Arctiidae Halysidota tessellaris JE Smith Bucculatricidae — Bucculatrix nr. albertiella Gelechiidae Trypanisma prudens Clemens Limacodidae Isa textula Harris Parasa indetermina Bdv Sabine stimulea Clemens Lymantriidae Dasychira obliquata Grt. & Rob. Noctuidae Acronicta increta Morr. Meganola miniscula Zeller Notodontidae Lochmaeus manteo Doubleday Natada gibbosa JE Smith Oecophoridae Machimia tentoriferella

Clemens

“Nomenclature and authors of Lepidoptera follow Hodges (1983).

Pseudotelphusa sp.) were preferentially occupying pre-existing (“old”) leaf ties. Because most leaf ties were colonized and maintained by a succession of larvae of various species, the local density of leaf ties formed early in the season should have positively in- fluenced the rate of colonization (and perhaps fo- livory) later in the season. The positive relationship between early season leaf tie density and the density of leaf ties formed over the entire season lent some support to this idea, but experimental studies are needed to separate innate host plant effects (some trees are simply more attractive to leaf tiers) from the domino effects arising from sequential occu- pancy of pre-existing ties.

The increase in abundance and species richness of insect herbivores with later date of leaf tie establish- ment (after controlling for tie age; Fig. 4) most likely reflected the seasonal increase in the size of the species pool of herbivores; more species were available to colonize a limited resource, resulting in more crowded leaf ties. There were a variety of late-season species not present ealier, whereas virtually all of the early summer species were also found in late summer, since most of these early species are bivoltine.

The rapid colonization of the artificial leaf ties by leaf tiers suggests that these species actively seek out pre-existing ties. Because many of these secondary oc- cupants were early instar caterpillars, it appears that female moths are selecting pre-existing shelters as oviposition sites (wherein hatching larvae typically take up residence). Ovipositing in pre-existing shelters may provide several benefits to developing offspring, in- cluding a favorable microclimate that decreases the risk of dessication, decreased construction costs (in

6

terms of both time and energy), and decreased ap- parency to visually foraging predators (Fukui 2001). There are also potential costs to occupying pre-existing shelters, most of which are related to the negative ef- fects of direct and indirect competition (Damman 1993). For example, pre-existing shelters are often highly damaged, contain large amounts of frass, and are often occupied by other caterpillars, all of which have been shown to increase the risk of predation by members of the third trophic level (Heinrich & Collins 1983, Steiner 1984, Mattiacci & Dicke 1995, Masashi 1999, Weiss 2003).

In addition, the surfaces of many “old” leaf ties have been skeletonized by previous occupants and would appear to offer little food resources to new colonists. However, because early instar caterpillars have rela- tively modest food requirements and later instars can add leaves to their natal shelter or create new shelters, it is quite possible that future food limitation has little bearing on adult oviposition decisions in this system. In this study, caterpillars colonized leaf ties of different ages in proportion to their relative abundance on the study trees, suggesting that ovipositing moths are not particularly selective about the age of leaf ties chosen for oviposition sites. Previous studies have shown that shelter-building caterpillars often choose leaves that promote effective and efficient shelter construction, even at the expense of food quality (Damman 1987, Hunter 1987, Reavey 1991, Loeffler 1996).

The diversity of ways in which caterpillars engineer their environment through the production of leaf shel- ters is only just beginning to be explored. More studies that investigate the costs and benefits of shelter-building as a putative adaptive trait are needed, as are studies that examine the ecological consequences of the behavior for plants and the communities of arthropods that associate with them. In addition, our understanding of the natural history of these small, inconspicuous, yet often abundant caterpillars is extremely limited; collecting and rearing efforts are needed to improve morphological descrip- tions and determine the host plant affinities and life his- tories of many of these poorly-known shelter-builders.

ACKNOWLEDGMENTS

I am grateful to R. Marquis, K. Boege, G. Chen, R. Forkner, R. Rios, K. Schultz, M. Weiss, and an anonymous reviewer for helpful comments on the manuscript; B. Schuette (Missouri Department of Natural Resources) for providing logistical support at the field site; D. Lill for field assistance; and the NSF (DEB-9700887), Sigma Xi, Web- ster Groves Nature Study Society and Trans World Airlines for funding.

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PRENTICE, R. M. 1965. Forest Lepidoptera of Canada. Vol. 4. Mi- crolepidoptera. Bulletin number 1142. Department of Forestry of Canada, Ottawa, Canada.

ReEAVEY, D. 1991. Do birch-feeding caterpillars make the right feeding choices? Oecologia 87:257-264.

STEINER, A. L. 1984. Olacmretiions on the possible use of habitat cues and token stimuli by caterpillar-hunting wasps: Eudynerus foraminatus (Hymenoptera: Eumenidae). Quaestiones Ento- mologicae 20:25-34.

Weiss, M. R. 2003. Good housekeeping: why do shelter-dwelling caterpillars fling their frass? Ecol. Letters 6:361-370.

Received for publication 30 January 2003; revised and accepted 25 July 2003.

Journal of the Lepidopterists’ Society 58(1), 2004, 7-12

A NEW SPECIES OF YPHTHIMOIDES (NYMPHALIDAE, SATYRINAE) FROM SOUTHEASTERN BRAZIL

ANDRE VICTOR LUCCI FREITAS

Museu de Hist6ria Natural and Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, Campinas, Sao Paulo 13083-970, Brazil

ABSTRACT. | This paper describes a new species of Yphthimoides (Nymphalidae, Satyrinae) from the highlands of the Serra do Cip6, Minas Gerais, southeastern Brazil. The position of this species is still uncertain within the genus, but it is near to Y. celmis (Godart) based on the male genitalia. Early stages are very similar to those of other species of Euptychiini. Adults are most active from 10:00 to 14:00 h, mostly on slopes pro- tected from direct winds. Males are territorial, and females wander through the area. Adults were observed feeding on six species of flowers.

Additional key words: Campos rupestres, Poaceae, Serra do Cip6, Yphthimoides cipoensis.

The Serra do Cipé is the southernmost part of a mountain system known as the “Serra do Espinhago,” extending through the center of Minas Gerais State, in SE Brazil. Most mae its area is included in the “Serra do Cipo National Park,” covering parts of the municipali- ties of Jaboticatubas, Santana ido Riacho, Morro do Pi- lar and Itambé do Mato Dentro. The region is mostly covered by open vegetation, especially rocky montane fields (“campo rupestre”) and Cerrado, with narrow galley forests bordering some rivers in the valleys (King 1956, Joly 1970, Moreira & Camelier 1977).

The region has many endemic species of plants and animals (Vanzolini 1982, Giulietti et al. 1987, Etero- vick & Sazima 2000, Sawaya & Sazima 2003), but the knowledge of insects including butterflies is minimal. Preliminary lists for the region show that it is espe- cially rich in endemic Hesperiidae and Lycaenidae; within the Nymphalidae, the Satyrinae are the group best represented (E. G. de Oliveira, in prep.). Any bi- ological information about this region is important, es- pecially considering the need for protection of the en- tire area.

The species described here was first collected by the author in 1996 near the Serra do Cipé National Park, and also observed in three subsequent visits to this area (1997, 2001, 2002). Attempts to identify this species indicated that it was an undescribed taxon.

The present paper describes this new species and illustrates the morphological characters of adults and immature stages.

MATERIALS AND METHODS

The species was studied in the Serra do Cipé, Mi- nas Gerais, SE Brazil. The mean annual temperature is 18°C (Nimer 1972, CETEC 1982). Annual rainfall averages 1600 mm, with a marked dry season from May to August. Observations were made in the “Juquinha” site, Santana do Riacho, Minas Gerais, on a hilltop with many rocky outcrops delimiting an area of about 300 m? where adults were usually abundant

(more than 60 marked in two days in 1996, and about 20-30 in 1997 and 2002). Adults were more common in the western section, where the slope was more pro- tected from winds.

Fertile eggs were obtained from two wild-captured females confined in plastic bags. Larvae were reared in plastic containers cleaned daily, with fresh plant mate- rial provided every two or three days (following Freitas 1991). Data were recorded on behavior and develop- ment times for all stages, and dry head capsules and pupal castings were kept in small glass vials. When there was Saffcient material, immatures were fixed in Kahle solution (AVLF collection). All measurements were made using a microscope fitted with a calibrated micrometric ocular. Egg size is presented as height and diameter, and head capsule size is the distance be- tween the most external ocelli (as in Freitas 1991). Taxonomic nomenclature follows Miller (1968) modi- fied by Harvey (1991), who treated the Satyrinae as a subfamily, downranking Miller’s subfamilies and tribes to tribes and subtribes, respectively. Nomenclature of wing veins follows Miller (1969), and of body setae fol- lows Hinton (1946).

Yphthimoides cipoensis Freitas, new species (Figs. 1-3)

Adults: Diagnosis. Eyes with minute sparse hairs, appearing naked without magnification: reddish brown with dark areas (varying in shape and position). Palpus length 1.5 times head height, brown with light brown hairs. Antenna (10-11 mm) extending to mid-costa; shaft dark brown, dorsally covered by dark brown scales, club with 16-17 segments, not conspicuously developed. Male wing venation shown in Fig. 2a. Forewing slightly elongated, hindwing outer margin wavy, especially in the males. Male foreleg with two elongated partially fused tarsomeres; fem: ale foreleg with five tarsomeres (Fig. 2c, d). The midleg and pal- pus are shown in Fig. 2b, e. Easily distinguished from other species of Yphthimoides by the lack of conspicu-

Fic. 1. Yphthimoides cipoensis from Serra do Cip6, Minas Gerais; adult male (top) and female (bottom). Left wings ventral, right wings dorsal.

ous ocelli on the wings and by the weakly marked lines crossing the wings: Fig. 3 compares Y. cipoensis with 15 other common species of Yphthimoides from Southern Brazil, showing that the general wing shape and color pattern of Y. cipoensis is quite different from those of the other species.

Description. Male (Fig. 1, top). Forewing length 21-22 mm; hind- wing length 15-18 mm (n = 10). Body entirely dark brown. Dorsal gr ann color dark brown with no markings, except for a thin orange- brown mar ginal stripe on the hindwing; covered with hairs, longer and more abundant on the basal portion, especially on the hindwing. Ventral ground color of wings same as dorsal; forewing crossed by a dark brown concave irre gular line extending from costa to CUI at two thirds from base, delimiting a distal area with scattered pale pink scales giving a bluish tint, and with a minute black dot in the space M1—M2 one fifth from wing margin; a dark brown zigzag sub- marginal line and an orange brown marginal line extending from costa to 2A. Hindwing with many scattered pale pink scales giving a bluish tint; a dark oun concave irregular line from costa to all margin, delimiting a distal area with bluish tint more pronounced; a series of five narrow white stripes over veins M1 to Cu2, the second (on M2) weaker; two minute black dots with white center, in spaces M1-—M2 and Cul—Cu2.

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Male genitalia unusual in shape within the Satyrinae (Fig. 2f-i). Uncus forming a dorso-ventrally flattened process, in the form of a wide spatula (Fig. 2g). Tegumen slightly bilobed; saccus very short and small. Gnathos appearing as two long pointed processes ‘almost the same size as uncus. Valvae elongated, ending in a single blunt point, with internal margin ornate bearing many well developed spines in three series: a basal rounded process with 6-8 small spines, an intermediate region with 2-3 long spines, and a terminal protu- berance with 3-7 small spines (number, size and disposition of spines in each series different on the two valvae of the same indi- vidual and among different individuals) (Fig. 2i). Aedeagus elon- gated with a conspicuous shelf on the ventral surface (Fig. 2h). Female (Fig. 1, bottom). Forewing length 22-24 mm; hindwing length 17-19 mm (n = 5). Body entirely ala brown. General oiler and pattern very similar to, but in general paler than that of males, with less bluish tint, and with the distal area paler than basal area on both wings.

Variation. Variation on the dorsal wing surfaces was absent in the sample studied, and differences in general darkness were in- versely correlated with wing wear. The ventral surface of the wings can be we »akly marked in some individuals of both sexes, and the white stripes on the hindwing usually disappear in old individuals. The ocelli can be absent in some individuals, and in one female an additional ocellus was seen in the space Cu2—1A in the hindwing.

Early stages: Egg (Fig. 4a). Spherical, beige, without ridges but with many small depressions in the nee Height 1.14-1.30 mm (mean = 1.197 mm, SD = 0.049, n = 7); diameter 1.14-. 24 mm (mean = 1.191 mm, SD = 0.04], n =7 Spaition 5-9 days (N = 30).

First instar (Fig. 4b-d). Head capsule black, with enlarged cha- lazae, bearing a pair of short scoli on vertex, each with two long nar- row setae ending in a fine point (Fig. 4b). Third stemmata larger than the other stemmata. Head capsule width 0.76-0.82 mm (mean = 0.789 mm, SD = 0.016, n = 25); scoli 0.08—0.12 mm (mean = 0.095 mm, SD = 0.010, n = 25). Body beige, smooth, with red longitudinal stripes; caudal filaments very short. Setae XD, D, SD and L thick- ened with clubbed tips; body chaetotaxy illustrated in Fig. 5. Maxi- mum length 7 mm. Duration 10-15 days.

Second instar. Head dark brown with two div erging short scoli on vertex. Head capsule width 1.14-1.30 mm (ccocetn = 1.213 mm, SD = 0.042, n = 19); scoli 0.16—0.20 mm (mean = 0.184 mm, SD = 0.014, n = 19). Body brown with many longitudinal stripes; caudal filaments more Maximum length 11 mm. Duration 6-13 days.

Third instar (Fig. 4e). Head medium brown, with two dorsal dark lines reaching bases of two diverging very short scoli on vertex. Head capsule width 1.84-2.00 mm (mean = 1.917 mm, SD = 0.051, n = 19); scoli 0.24-0.30 mm (mean = 0.264 mm, SD = 0.021, n =19). Body brown with many longitudinal stripes; caudal filaments short. Maximum length 17 mm. Duration 7-10 days.

Fourth (last) instar (Fig. 4f, g). Head lighter brown, with two dorsal dark lines to bases of div erging short scoli on vertex, integrat- ing head and body stripes. Head capsule width 2.81-3.39 mm (mean = 3.101 mm, SD = 0.190, n = 10); scoli 0.39-0.43 mm (mean = 0.409 mm, SD = 0.021, n = 10). Body brown with many longitudinal stripes; one dorsal stripe conspicuously dark; ventral region dark brown; legs and prolegs light brown; caudal filaments drank Maxi- mum length 30 mm. Duration 17-20 days.

Pupa ( (Fig. 4h-j). Short and smooth; mostly beige, with short squared ocular caps; cremaster dark in ventral portion; dorsal ab- domen with a paired series of short subdorsal white protuberances; alar caps light at the edge, with lightened visible tracheae. Total length 12— 13 mm. Duration 20-25 days.

Etymology. The specific name refers to the distribution of this species, which apparently is restricted to the Serra do Cipo.

Holotype: adult male from “Juquinha” (19°15’S, 43°33W), 1370 m, Serra do Cip6, Santana do Riacho, Minas Gerais, south- easter Brazil, collected by A. V. L. Freitas on May 6, 1996. In the collection of the Departamento de Zoologia, Universidade Federal do Parana, Curitiba, Parana, Brazil (collection reference number: DZ 5.132). Labels on the holotype (three labels, separated by trans- verse bars): /HOLOTIPO/ SERRA DO CIPO, JUQUINHA, SAN-

VOLUME 58, NUMBER |

R3 R4 R2 R5 R1 M1 sc M2 M3 Cut Cu2 2A

Sc+R1

Rs M1 h M2 M3 2A Cut

Cu2 1A

par Ue

Fic. 2. Morphological characters of Yphthimoides cipoensis. a, Male wing venation, hindwing above and forewing below; b, Male midleg:

c, Male foreleg; d, Female foreleg; e, Male palpus; f, Lateral view of male genitalia; g, Dorsal view of tegumen and uncus; h, Aedeagus (lateral above, ventral below); i, Right valva upper view (external above, internal below).

TANA DO RIACHO, MG, BRASIL, 19°15’S, 43°33’W, 1370 m, 6-V-

GB, SIL GRIST! Goce SPEED Distribution. This species is very local, and was 396, A.V. L. FREITAS leg. 69.132, 6 0 2 1 arog Paratypes: Three adult males and one adult female, same site as observed in only three sites in the Serra do Cipo. In holotype, collected in May 2002, in the collection of the author.

addition to the population of “Juquinha,” a second

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Fic. 3. Comparison of Yphthimoides cipoensis with another 15 species of Yphthimoides of Southern Brazil. Top to bottom, left to right; Y.

cipoensis Freitas, Serra do Cip6, Minas Gerais; Y. celmis (Godart), Viamao, Rio Grande do Sul; Y. renata (Stoll), Mogi Guagu, Sao Paulo; Y. straminea (Butler) (probably a dry season form of Y. renata), Luiz Anténio, Sa0 Paulo; Yphthimoides sp., Campinas, Sao Paulo; Y. affinis (But- ler), Campinas, Sao Paulo; Y. ochracea (Butler), Campos do Jordao, Sao Paulo; Y. yphthima (C. & R. Felder), Campinas, Sao Paulo; Y. borasta (Schaus), Igarata, Sao Paulo; Y. grimon (Godart), S40 Sebastiao, Sao Paulo; Yphthimoides sp., Teodoro Sampaio, Sao Paulo; Y. manasses (C. & R. Felder), Itirapina, Sao Paulo; Yphthimoides sp., Sao Sebastiao, S40 Paulo; Yphthimoides sp., Cotia, S40 Paulo; Y. viviana (Romieux), Santa

Barbara, Minas Gerais; Yphthimoides sp. (ca. castrensis), Maquiné, Rio Grande do Sul.

population was found by Marcio Uehara-Prado in “Travessao” (19°20’S, 43°31’W, elevation 1100 m), in July-August 2001, and W. W. Benson found a third population in a private area not far from the “Serrote” (19°17’S, 43°33’W, elevation 1200 m) in May 2002. The species probably occurs in other similar habitats in the region.

Behavior and Natural History. Oviposition be- havior was not observed, and the host plant in the field is unknown. In the laboratory, larvae easily accepted Goosegrass Eleusine indica (L.) Gaertn. (Poaceae), a common introduced grass in Brazil. In plastic bags, eggs were usually laid singly on the plastic surface (suggesting that in nature eggs are also laid singly). Adult males are territorial, while females wander through the landscape (W. W. Benson & AVLF unpub- lished data). Adults were observed feeding on flowers

of Declieuxia sp. (Rubiaceae), Leucothoe sp. (Eri- caceae), Hyptis sp. (Lamiaceae), Piptolepis buxoides Schultz-Bip and two unidentified species of Vernonia (Asteraceae).

DISCUSSION

The genus Yphthimoides was erected by Forster (1964) to include about 15 species of medium-sized predominantly brown Neotropical Satyrinae species. Because the diagnosis given by Forster was vague, about 22 species have been included in this genus un- til now (G. Lamas pers. com.). Preliminary studies on the immatures (AVLF unpublished results) suggest that this genus is an unnatural assemblage of several distinct groups (in the description of Yphthimoides, Forster already recognized two distinct groups of species), and needs a major reorganization into five or

VOLUME 58, NUMBER 1

Sed wom me es ST SPYE Pare peer

ree SM Atoen atom asoe.

Fic. 4. Early stages of Yphthimoides cipoensis. a, Egg; b, First instar head capsule; ec, d, First instar (dorsal, lateral); e, Third instar (dorsal): f, g, Last instar (lateral, two larvae dorsal): h, i, j, Pupa ( (Gorelt ventral, lateral).

more genera. The tentative placement of Y. cipoensis

in Yphthimoides was based on the paper by Forster

(1964), and by the similarity of the male genitalia with

iY caine (Godart) (male genitalia figured in Forster

1964:100), a species very different from Y. cipoensis

(Fig. 3). However, this classification may require a re- assessment once our knowledge of Yphthimoides sys- tematics is improved.

Based on current information, this species is con- sidered endemic to the Serra do Cip6 region, and dis-

\, alk, -

Va T2

\7

T3 Ai A2/A3-6 A7 A& AQ A10

legs proleg anal proleg

Fic. 5. Chaetotaxy of the first instar larva of Yphthimoides cipoensis.

tributed in few scattered local colonies. Like many other species endemic to this mountain system, it may be considered vulnerable to extinction due to habitat destruction. Only one colony is known from the area inside the National Park, and the colonies outside are threatened by destruction of surrounding habitat, fire and uncontrolled eco-tourism. Additional efforts to discover new colonies of this species and population studies of the adults could be important to help in the preservation of the species and of the whole Serra do Cip6 system.

ACKNOWLEDGMENTS

This study was conducted as part of a Post-Doctoral project on Satyrinae (Nymphalidae) ecology and systematics (BIOTA-FAPESP program, grants 98/05101-8 and 00/01484-1), and as part of the ac- tivities of the undergraduate course of animal ecology (BE-780) of the Unicamp. This study was supported in part by the National Sci- ence Foundation (DEB-0316505). Keith S. Brown Jr. helped with pictures of immatures. Jorge Tamashiro identified the plant species. Thanks to Keith S. Brown Jr., J. Vasconcellos-Neto, W. W. Benson, Gerardo Lamas, Lee D. Miller, J. Y. Miller, and Carla Penz for helping in diverse phases of the development of the manuscript. I would like to thank the direction of the Parque Nacional da Serra do Cip6, Minas Gerais, for logistic support during field work.

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

LITERATURE CITED

CETEC. 1982. Principais atividades realizadas para a implantagao do Parque Estadual da Serra do Cipé. Relatério técnico. Belo Horizonte, MG.

ETEROVICK, P. C. & I. SAziMa. 2000. Structure of an anuran commu- nity in a montane meadow in southeastern Brazil: effect of sea- sonality, habitat, and predation. Amphibia-Reptilia 21:439-469.

Forster, W. 1964. Beitriige zur Kenntnis der Insektenfauna Bo- liviens XIX. Lepidoptera III. Satyridae. Veréffentlichungen der zoologischen Staatssammlung Miinchen 8:51-188, pls. 27-35.

Freitas, A. V. L. 1991. Variagao morfolégica, ciclo de vida e sis- tematica de Tegosa claudina (Eschscholtz) (Lepidoptera, Nymphalidae, Melitaeinae) no Estado de Sao Paulo, Brasil. Rev. bras. ee 35:301-306.

GruLieTtl, A. M., N. L. MENEZES, J. R. PrrANI, M. MEGURO & M. G.L. ae 1987. Flora da Serra do Cipé: caracteriza- cao e lista das espécies. Boletim de Botanica da Universidade de Sao Paulo 9:1-151

Harvey, D. J. 1991. Higher classification of the Nymphalidae (Ap- pendix B). Jn Nijhout, H. F., The development and evolution of butterfly wing patterns. Smithsonian Press. Pp. 255-273.

HinTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Trans.R. ent. Soc. London 97:1-37.

Joty, A. B. 1970. Conhega a vegetagao brasileira. Poligono, Sao Paulo, Sao Paulo, Brasil.

Kinc, L. C. 1956. A Geomorfologia do Brasil Oriental. Rev. Bras. Geogr. 18:147-265

MILLER, L. D. 1968. The higher classification, phylogeny and zoo- geography of the Satyridae (Lepidoptera). Mem. Am. Entomol. Soc. 24:iii + 174 pp.

. 1969. Nomenclature of wing veins and cells. J. Res. Lepid. 8(2):37-48.

Moreira, A. A. N. & C. CAMELIER. 1977. Geomorfologia do Brasil—Regiao Sudeste. Rio de Janeiro, IBGE vol. 3:1-50. NIMER, E. 1972. Climatologia da Regiao sudeste do Brasil. Intro- ducao a climatologia dinamica. Rev. Bras. Geogr. 34:3-48. Sawaya, R. J. & I. Sazima. 2003. A new species of Tantilla (Ser- pentes: Colubridae) from southeastern Brazil. Herpetologica

59(1):119-126.

VANZOLINI, P. E. 1982. A new Gymnodactylus from Minas Gerais, Brasil, with remarks on the genus, on the area and on montane endemisms in Brazil. Papéis Avulsos de Zoologia 34:403-413.

Received for publication 07 March 2003; revised and accepted 25 July 2003.

Journal of the Lepidopterists’ Society 58(1), 2004, 13-20

SPHINX MOTH POLLINATORS FOR THE ENDANGERED WESTERN PRAIRIE FRINGED ORCHID, PLATANTHERA PRAECLARA IN MANITOBA, CANADA

A. RICHARD WESTWOOD! AND CHRISTIE L. BORKOWSKY Department of Biology, University of Winnipeg, Winnipeg, Manitoba R3B 2E9, Canada

ABSTRACT. The westem prairie fringed orchid, Platanthera praeclara (Sheviak & Bowles), is an endangered species in North America. In Manitoba orchids produce lower numbers of seed capsules than more southern populations. Exploration of the pollination biology for P. praeclara is critical to preserve this endangered species. This study identified the pollinators of P. praeclara using cone and malaise traps and tested the effectiveness of marking pollinators with traces of Day-Glow® orange marker powder. Although Lepidoptera were numerous in or- chid plots during daily observation periods, including day flying Sphingidae, none were pollinators for P. praeclara. Among the 5856 insects from 49 families captured over 45 trapping days, six sphinx moths, two specimens of Hyles gallii (Rottenburg) and four specimens of Sphinx drupifer- arum J.E. Smith (Sphingidae), were found with two or more orchid pollinia attached to their eyes and were confirmed as pollinators of P. praeclara. S. drupiferarum is uncommon in southern Manitoba and H. gallii appears to be a less efficient pollinator than S. drupiferarum. Pro-

boscis length, eye width and flight period may influence the efficiency of biotic pollination of the orchid.

Additional key words: Sphingidae, pollinia, pollination, P. praeclara, S. drupiferarum, H. gallii.

The western prairie fringed orchid, Platanthera

praeclara (Sheviak & Bowles 1986), occurs in areas of remnant tall grass prairie in southeastern Manitoba. Prior to its discovery in the mid 1980's near Tolsoi, Man- itoba, P. praeclara was not known to exist in Canada (Johnson 1985, 1991). The Manitoba population, at its maximum has consisted of approximately 21,000 indi- vidual plants (although the population may fluctuate widely from year to year) and it is the largest of four metapopulations (over 300 plants) in North America. The remaining populations occur in North Dakota, South Dakota, Minnesota, Kansas, Nebraska, and Iowa (Sheviak & Bowles 1986, Bjugstad & Fortune 1989, Bray & Wilson 1992, Bjugstad-Porter 1993, Pleasants & Moe 1993, Davis 1994, Sieg & King 1995, US. Fish & Wildlife Service 1996, Hof et al. 1999).

The orchid is protected under Manitoba’s Endan- gered Species Act and has been placed on the endan- gered species list in both Canada and United States (Collicutt 1993, Davis 1995). In Manitoba, P. praeclara often has low fruit set and subsequent seed production (Borkowsky 1998). Although the western prairie fringed orchid will produce a vegetative form, there is little evidence that vegetative ngpractiaition occurs in P. praeclara (Bowles 1983, Sather 1991, Sieg & King 1995, Hof et al. 1999). A plant may also go ere as is typical in other species of orchids (Nilsson 1992). Therefore, the recruitment of new plants is dependant primarily upon successful pollination and subsequent seed production. In surveys of over 1000 plants Borkowsky (1998) found that only 2.1% of orchid stems produced one or more seed capsules annually in Manitoba between 1994 and 1998. In more southern orchid populations the percent of stems that produce

' Corresponding author: R. Westwood, email: r.westwood@ uwinnipeg.ca; Phone (204)-786-9053; Fax: (204)-774-4134.

seed capsules can range up to 49% (Bowles 1983, She- viak & Bowles 1986, Cuthrell 1994). The authors have hypothesized that reduced seedpod production may be linked to low pollination success in Manitoba. The level of pollination success for P. praeclara and the identity of the orchid pollinators are unknown in Manitoba.

Pollination is the process in which pollen grains are transferred to the stigma, which is followed by fertil- ization of the ovules aid development of seeds (Proc- tor et al. 1996). Many orchids require a biotic organ- ism (a pollination agent or pollen vector) to transport the pollen to the stigma (van der Pijl & Dodson 1966, Faegri & van der Pijl 1979). In P. praeclara the most striking visual characteristic of the flower is a large, deeply fringed, tri-lobed lower lip and a long, slender spur containing nectar that suspends backward from the flower (Figs. 1, 2). The nectar spur may be 36-55 mm long with a maximum diameter of 2.7 + 0.5 mm (Sheviak & Bowles 1986). Orchids grow to 38—S5 cm tall (Sheviak & Bowles 1986) with the determinant in- florescence containing 7 to 12 flowers (Sheviak & Bowles 1986, Pleasants 1993). The pollinium (Fig. 1) is a specialized structure that consists of pollen, a col- umn and viscidium (Nilsson 1992). In P. praeclara the minute grains of pollen are arranged into subunits called massulae (Pleasants & Moe 1993). These sub- units form a bi-lobed mass that is attached to the col- umn, which is secured to the viscidium (the entire structure is termed the pollinium). The pollinium is sheathed, with the exception of the viscidium, which is exposed and adapted to cement itself to the pollinator (Bowles 1983). Each flower has one pollinium located on either side of the stigmatic surface. This allows for a 6 to 7 mm separation between each viscidium (She- viak & Bowles 1986). The opening to the nectar spur is located immediately below the stigmatic surface.

The small opening to the nectar spur restricts the

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Fic. 1. Platanthera praeclara flower, anterior view, showing pollinium (a), and the opening to nectar spur (b). 8

position of a potential nectar seeking insect, and in- creases the likelihood that one or both viscidia will come into contact with a likely pollinator. The pollinium is pulled from the sheath when a pollinator contacts the viscidium and then withdraws from the flower (Darwin 1904, Dressler 1981, 1993).

Orchids also produce a scent or fragrance which may be weak or absent in the daytime, increases in in- tensity at dusk and may remain strong until sunrise de- pending on the age and condition of the flower (She- viak & Bowles 1986). The relationship between the attractiveness of the fragrance to pollinators and the intensity of the scent is unknown in P. praeclara.

To be identified as a pollinator and to rule out in- discriminate visitors to the flower, the organism must make regular visits to the flowers during its lifetime and effectively deposit the pollen on the stigma (Faegri & van der Pijl 1979). The flower—pollinator relation- ship may also be controlled by pollinator behavior, mouthpart morphology or taxonomy and by flower morphology (Faegri & van der Pijl 1979, Wyatt 1983). Wyatt (1983) identified nine forms of biotic pollination in orchids including sphingophily (Sphingidae—hawk- moths), phalaenophily (small moths), psycophily (but- terflies), melittophily (bees), myophily (syrphid and bee flies), sapromyophily (carrion and dung flies), can- tharophily (beetles), ornithophily (birds) and chi- ropterophily (bats).

The potential pollinators for P. praeclara in Mani- toba are not known. Although the literature suggests that night-flying members of the Sphingidae may be key pollinators of P. praeclara, other members of this orchid genus and several closely related genera may be pollinated by butterflies, other moth species, certain Diptera, Coleoptera or Hymenoptera (van der Pijl &

Fic. 2. Platanthera praeclara flower, lateral view, showing nec- tar spur (a).

Dodson 1966, Patt et al. 1989, Robertson & Wyatt 1990, Bowles et al. 1992, Larson 1992).

The floral characteristics of P, praeclara indicate a moth pollination method (Faegri & van der Pijl 1979, Sheviak & Bowles 1986, Luyt & Johnson 2001), most likely sphingophily or phalaenophily. P. praeclara has no developed landing platform on the flowers, there- fore hovering pollinators are favored and butterflies may be excluded (van der Pijl & Dodson 1966, Dressler 1981). To retrieve nectar, the insect must cor- rectly align itself with the flower as it inserts its pro- boscis, which increases the likelihood that the pollina- tor will contact one or both of the viscidia which will adhere to the proboscis or eyes of the insect (Sheviak & Bowles 1986). Considering the length of the nectar spur of P. praeclara and position of the viscidia, the list of potential pollinators is further reduced to Lepi- doptera with a very long proboscis (e.g., moths belong- ing to the Sphingidae or hawkmoth family). Few, if any, observations of pollination of fringed orchids by hawkmoths have been made in the field (Bowles 1983, Sheviak & Bowles 1986, Pleasants & Moe 1993). Based on proboscis length, Sheviak & Bowles (1986) suggested that the following species of moths could be potential pollinators of P praeclara in the United States: Eumorpha achemon (Drury), Hyles lineata (F.), Sphinx drupiferarum J.E. Smith and Sphinx kalmiae J.E. Smith. Cuthrell (1994) collected two specimens, one each of Sphinx drupiferarum and Eumorpha achemon with viscidia from P. praeclara attached to the head, from a light trap adjacent to a field of orchids in the United States.

To better understand the pollination biology of P. praeclara in Manitoba diurnally active Lepidoptera including day flying Sphingidae, nocturnally active

VOLUME 58, NUMBER 1

Lepidoptera and other insect Orders were surveyed as potential pollinators of P. praeclara. Several types of passive traps were tested to capture P. praeclara polli- nators, and a method to mark individual pollinators was investigated.

STUDY SITE AND METHODS

Five field experiments were established between 1997 and 1999 to determine the identity of P. praeclara pollinators. Three study plots were located in the Manitoba Tall Grass Prairie Preserve, near Tolsoi, Manitoba (49°05’N, 96°49’W) and two additional moth light trapping sites were located near Lonesand, Manitoba (approximately 21 km east of the Preserve) and near Grunthal, Manitoba (approximately 26 km northwest). The climate is a boreal continental regime with mean temperatures of 19.6°C and —18.8°C for July and January, respectively. Fifty-five percent of the annual precipitation (mean 579.1 mm) falls during the period of May through August (Canadian Climate Pro- gram 1993). Drainage in much of the prairie is poor with soil composed of lacustrine parent material, sandy loam to clay loam upper horizons and a thin organic surface layer. Stones, rocks and occasionally boulders are also scattered across the prairie (Canada Soil In- ventory 1989, Moore & Fortney 1994).

The Tall Grass Prairie Preserve has an abundance of grasses, including Big bluestem (Andropogon ger- ond Vitman) and Little bluestem (Schizachyrium sco- parium (Michx.) Nash) (Gramineae) and is inter- spersed with bluffs of various deciduous tree species, including willow (Salix spp.) and poplar (Populus spp.) (Henne & Diehl 2002).

Three 50 x 50 m plots were established in fields were orchids were numerous. Each plot contained a minimum of 25 orchids. Plots were separated by a dis- tance of 500 to 1000 m. To establish that there were no day flying insects pollinating the orchids, 15 individual orchids were tagged in each of the three plots (total of 45 plants) in 1997 and 1998. Each orchid was visually observed for 15 minutes on at least three separate days during the flowering period in 1997 and 1998 between 1100 i to 1500 h to determine if insects contacted plants and were responsible for removal of one or more of the pollinium from the flowers, either by acci- dent or for the purpose of obtaining nectar or pollen. The number and identity of potential pollinators (Or- der, Family and Genus/species if known) landing or crawling on the orchids was recorded.

Inverted cone insect traps were used to sample po- tential pollinators from individual orchids. Cone traps were constructed from light gauge steel tubing and wire and covered in fine mesh screen. Cone traps were

approximately 40 cm in length and 25 cm in diameter. An inverted wire mesh cone was placed at the base of the trap (much like a minnow trap) and a hinged trap door placed on the top of the trap. Three legs attached to the bottom of trap allowed it to be placed over an individual orchid. The legs were of sufficient length that they could be pushed into the ground around the orchid, allowing the mouth of the inverted cone to be suspended approximately 5 to 10 cm above the termi- nal flower of the orchid. Traps were emptied daily be- tween 0630 h and 0830 h and again between 1900 h and 2030 h. Captured insects were identified and ex- amined for presence of orchid pollinia. One cone trap was placed in each of the three plots and rotated be- tween plants during the flowering period. Cone traps were placed over orchids that had the most flowers open with intact pollinia available. Traps remained in place over an individual orchid for 24-48 hours and were placed in the plots between | July and 15 July in 1997, 1998 and 1999.

Malaise insect traps (Bioquip® Equipment Special- ities) were to used to sample for potential pollinators over a groups of orchids (generally five or more indi- vidual plants). Each trap was approximately 2 m in height with a glass container at the top to collect trapped insects. A piece of Vapona® insecticide strip (0.2 cm?) was placed at the end of the collecting cylin- der to kill captured insects. One malaise insect trap was placed within each of the three plots and rotated among groups of orchids during the flowering period. Traps were emptied daily between 0630 h and 0830 h and again between 1900 h and 2030 h. Insects were identified and examined for presence of orchid pollinia. Traps were placed in the field between 1 July and 15 July in 1997, 1998 and 1999. Traps were ro- tated within the plot to new groups of orchids every three to five days.

In 1999, five orchids in each plot under a malaise trap or one orchid under a cone trap were chosen to test the effectiveness of Day-Glow® orange marker dye powder in identifying g potential pollinators. A small amount of Day elon orange marker dye powder was applied with a extra fine nylon brush to the centre of each flower of the orchids chosen. Insects captured in the traps were identified and examined for presence of pollinia removed from the orchids and also exam- ined in a dark room under an ultra-violet light for the presence of the Day-Glow® powder. Insects with marker particles adhering to their bodies were consid- ered to have come in contact with the treated flowers.

To determine moth flight periods, two Wards all- weather insect traps® (Wards Natural Science) were placed approximately 1 km to the west of the orchid

16

plots and 5 km to the south of the plots. Two additional

Vard’s all-weather insect traps® were located at Lone- sand and Grunthal, Manitoba to augment moth cap- tures at the Tall Grass Prairie Preserve. Traps were op- erated from 1 May to 31 August, 1997-1999. Traps had an eight watt ultra-violet fluorescent bulb as an at- tractant. The traps were used to survey nocturnal Lep- idoptera and determine the flight periods of potential orchid pollinators. Flight periods were considered to include the period between date of first and last cap- ture in each year. Traps in the Tall Grass Prairie Pre- serve were placed in open areas surrounded by mature trees and were not visible from orchid plots. Lepi- doptera captured in traps were sorted, pinned and identified to species.

The flowering period for orchids in plots was recorded during the study. Flowering period was de- fined as the period between the appearance of the first flower and last flower in a plot and peak flowering date was defined as the date when the most flowers were fully open in a plot. The mean overlap in days between moth species flight period and orchid flowering period for all years was calculated. Mean overlap flight period data was square root transformed to satisfy assump- tions of normality and heterogeneity of variance for analysis of variance (ANOVA) (SPSS Inc. 1999). Where ANOVA was significant a least-significant dif- ference (LSD) test was used to determine the differ- ences between means (@ = 0.05) because of its consis- tency (Saville 1990). To determine if potential pollinators would fit pollinia distance separation re- quirements and nectar spur depth requirements of P. praeclara, measurements were made on a minimum of five individuals from all sphinx moth species collected in the various trap types. Pinned moths were softened in a relaxing chamber and the length of the proboscis, the distance between the outer margins of the com- pound eyes and distance between the inner margins of compound eyes where measured in mm under a dis- secting microscope. For data analysis the proboscis length was square root transfor med and distance be- tween outer eye edges and distance between inner eye edges were log tector med to satisfy assumptions of normality and theexomenaitey of variance. Morphologi- cal measurements hetween species were subject to ANOVA and where ANOVA was significant differ- ences between means were identified using a LSD test. To examine the relationship between proboscis length and ability of pollinators to retrieve nectar, the length of nectar spurs and the depth of nectar within the spur (distance from distal end of spur to top of nectar line) was measured for orchids in each of the three plots.

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TABLE 1. Number and identity of insect Orders and Families collected in cone and malaise traps 1997-1999.

Order Family/Subfamily n Ephemeroptera Baetidae i Heptageniidae il Odonata Coenagrionidae 3 Orthoptera Acrididae 5 Plecoptera Perlidae 1 Hemiptera Miridae 21 Reduviidae 2 Pentatomidae 2 Homoptera Cercopidae 5 Cicadellidae 13 Neuroptera M antispidae 2 Chrysopidae 9 Coleoptera Carabidae 8 Scarabaeidae 5 Elateridae 2 Lampyridae 12 Cleridae 5 Coccinellidae 8 Tenebrionidae 1 Chrysomelidae 5 Curculionidae 3 Mecoptera Panorpidae 2 Trichoptera Limnephilidae 6 Diptera Tipulidae 26 Culicidae 55 Chironomidae 24 Simuliidae 15 Tabanidae 4450 Syrphidae 45 Muscidae 546 Calliphoridae 228 Sarcophagidae ll Lepidoptera Pyralidae 14 Pterophoridae 2 Tortricidae 34 Gelechiidae 40 Geometridae 18 Arctiidae 3 Noctuidae 34 Lasiocampidae 15 Sphingidae 6 Hesperiidae 10 Nymphalidae 5 Satyridae 2 Hymenoptera Ichneumonidae 23 Vespidae 2 Sphecidae 3 Megachilidae 2 Apidae 19

Identification of insects was based upon Hodges (1971), Rockburne and Lafontaine (1976), Morris (1980), Hodges et al. (1983), Covell (1984), Borror et al. (1989), Klassen et al. (1989), Layberry et al. (1998) and Handfield (1999).

RESULTS

Observational survey. In 1997 and 1998 orchids were observed between 1100 h and 1500 h for approx- imately 70 hours. Although numerous insects (many

VOLUME 58, NUMBER 1

TABLE 2. Summary of six Sphingidae collected from cone and malaise traps with attached pollinia 1997-1999.

# of Pollinia Marker

Date Species Type pollinia location powder 11 July 1997 Hyles gallii cone 2 head n/a 1] July 1997 Sphinx drupiferarum malaise 5 head n/a 15 July 1998 Sphinx drupiferarum malaise 11 head n/a 15 July 1998 Sphinx drupiferarum malaise 3 head n/a 20 July 1998 Sphinx drupiferarum cone 7 head n/a

6 July 1999 Hyles gallii cone 2 head present

belonging to pollinating families or genera) frequented the plots during the daily observation periods no indi- viduals were observed to seek nectar or pollen from the orchids, to use the flowers as a resting platform or to sun themselves on the orchids. Individuals of the sphinx moths Hemaris thysbe (Fabricus) (16 individuals) and Hemaris diffinis (Boisduval) (28 individuals) were observed in the orchid plots during the observation peri- ods but they were not attracted to orchids, despite re- peatedly visiting nearby flowering herbs for nectar.

Trapping experiments. Between 1997 and 1999 the cone and malaise traps caught 5856 individual in- sects from 49 families over 45 trapping days (Table 1). The only insects found to have pollinia attached to their bodies belonged to the family Sphingidae (Table 2). Six sphingid moths, two specimens of the Bedstraw hawkmoth, Hyles gallii (Rottenburg), and four speci- mens of the Wild cherry sphinx, S. drupiferarum were collected with two or more pollinia attached to the eyes (Figs. 3, 4). H. gallit had 2 pollinia per moth, while S. drupiferarum had 3 to 11 pollinia per moth.

Marking experiment. In 1999 one sphinx moth, H. gallii, was collected from a cone trap with traces of Day-Glow® orange marker dye powder on both eyes and the proboscis. Both pollinia attached to the eyes also had traces of powder, primarily on the massulae and viscidium.

ley

Fic. 3. Anterior/lateral view, Sphinx drupiferarum J. E. Smith with pollinia from Platanthera praeclara attached to eyes.

Flight period. The flight periods for sphingid species found in the vicinity of the Tall Grass Prairie Preserve were based on the catches from the four black light traps during the period of 1997 to 1999 (Table 3). Flowering dates for P. praeclara are also shown in Table 3. Generally the moths collected were most abundant in the first several weeks of the orchid flowering period. The flight periods of S. drupiferarum and H. gallii overlapped with orchid flowering by 34.6% and 45.3% respectively.

Morphological measurements. The proboscis length, width across the eyes and distance between the inner eye margins were measured for the 15 species of Sphingidae collected in the vicinity of the Tall Grass Prairie Preserve (Table 4). The mean orchid nectar spur length was 45.27 mm (n = 1016, SE = 0.134) and the mean depth of nectar within the spur was 12.44 mm (n = 1016, SE = 0.201).

DISCUSSION

The results confirm observations by Faegri and van der Pijl (1979), Sheviak and Bowles (1986), Cuthrell (1994) and Luyt and Johnson (2001) that P. praeclara is pollinated by nocturnal Lepidoptera confined to the family Sphingidae. It appears that there are no diur- nally active insects that seek nectar or pollen from P.

Fic. 4. Anterior/lateral view, Hyles gallit (Rottenburg), with pollinia from Platanthera praeclara attached to eyes. Note the large viscidium cemented to lower-center of eye, with massulae projected forward.

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TABLE 3. Flight period dates, orchid flowering periods, number of individual sphinx moths collected in black light traps and percent overlap between flight and flowering dates in the vicinity of the Tall Grass Prairie Preserve and for orchids 1997-1999.

Peak orchid flowering date Orchid flowering dates Sphingidae—flight period dates Ceratomia undulosa Harris Sphinx chersis (Hubner)

Sphinx kalmiae ].E. Smith Sphinx luscitiosa Clemens Sphinx drupiferarum J.E. Smith Smerinthus cerisyi Kirby Smerinthus jamaicensis (Drury) Poanes excaecatus (J.E. Smith) Poanes myops (J.E. Smith) Cressonia juglandis (J.E. Smith) Pachysphinx modesta (Harris) Hemaris thysbe (Fabricus)* Hemaris diffinis (Boisduval)* Darapsa myron (Cramer)

Hyles gallii (Rottenburg)

% Overlap of flight

Dats period & flowering 1997 1998 1999 n! period days + SE 2 July 4 July 7 July 22 June-16 July 19 June-20 July 23 June-19 July 12 June-12 July 7 June-18 July 9 June-31 July 14 92.0 + 4.9c° 31 May-2 July 11 July absent 6 14.1 + 13.9a 1-26 July 5-24 June 10 June-18 July 8 58.0 + 23.2abe absent 20 May—30 June 19 June-11 July rf 34.6 + 19.9abe 11 July 1 June-25 July 24 June 5 34.6 + 32.7abe 28 May-27 June 21 May-29 July 16 May-25 July 58 73.7 + 26.3be 25 May—30 June 1 June-10 July 29 May-18 July 123 65.3 + 18.2be 3 June-20 July 11 June-15 July 5 June-20 July 37 94.6 + 9.2c 11-28 June 9 June—4 July 1-27 June 34 29.3 + 16.9abe absent 8-30 June absent 5 11.7 + 11.6a 27 May-28 June 1] June-5 Aug 9 June-9 July 61 63.0 + 20.8be 31 May-25 June 5-28 June 4 June-2 July 16 25.3 + 6.9abe 21 May-2 July 10 June-10 July 1 June-13 July 8 62.0 + 10.4be absent 15 May-6 July absent 5 18.0 + 16.0a 15 June-31 July 31 May-29 June 6 July 18 45.3 + 28.5abe p = 0.040 F,, ,, = 2.086

‘Number of moths collected 1997, 1998 & 1999.

* Means within each column followed by the same letter are not significantly different (Fisher's LSD, p > 0.05). 34H. thybe and H. diffinis collected by sweep net in the vicinity of the Tall Grass Prairie Preserve.

praeclara. The presence of pollinia on one H. gallii specimen and four S. drupiferarum specimens and the marker powder and pollinia on another specimen of H. gallit confirm these species as pollinating agents of P. praeclara in Manitoba. Despite the larger size of the malaise trap covering more orchids, there was little difference in the capture rates of moths with pollinia between trap types. The Day-Glow® orange marker dye powder successfully mar ked one pollinator.

Pollinia were attached to the center of each eye on both moth species. $. drupiferarum had more pollinia per specimen. The distance between the outer edges of the eyes was greater in S. drupiferarum than H. gal- lit (Table 4), but it is unknown if this small difference in eye separation (0.43 mm) would affect the attach- ment of pollinia. Perhaps S. drupiferarum is more ag- gressive at retrieving nectar and presses harder into the centre of the orchid thus capturing more pollinia. Alternately H. gallii may visit the orchids less fre- quently peeiene in fewer opportunities to remove pollinia, or be deconmmed from visiting orchids once several pollinia have been attached to the eyes. H. gal- lit is more numerous in the vicinity of the orchids than S. drupiferarum, thus it seems likely that they are less attracted to the orchids.

Cuthrell (1994) described two specimens of S. drupiferarum (one caught by light trap and one a mu- seum specimen) and one specimen of E. achemon

(Drury) (collected by light trap) with P. praeclara pollinia attached to the eyes. The present study adds H. gallii to this list of pollinators while E. achemon does not occur in Manitoba.

The mean proboscis lengths for S. drupiferarum and H. gallii captured in this study were 38.40 mm and 33.50 mm, respectively. Subtracting the mean nectar depth from total nectar spur length provides a distance of 32.83 mm, thus it appears that a proboscis length of 30-35 mm is required to reach the nectar in P. praeclara. Based on proboscis length, Sphinx chersis (Hubner) and S. kalmiae may also be potential pollina- tors in Manitoba (Table 4). These species had a signif- icantly longer proboscis than all other species col- lected (Table 4), with no other species having a mean proboscis length greater than 23 mm. The separation of inner and outer eye margins between species was less distinct with some species having a short proboscis yet still having similar eye positioning and separation (Table 4).

Orchid flowering periods and moth flight periods are restricted in terms of overlap in Manitoba. The overlap in more southern areas of the range of P. praeclara may be greater and therefore higher levels of pollination may occur resulting in more seed cap- sules per plant. S. drupiferarum is uncommon in southern Manitoba and the most numerous sphingid species in the Tall Grass Prairie Preserve do not possess

VOLUME 58, NUMBER 1

TaBLE 4. Length of proboscis, the distance between the outer margins of the compound eyes and distance between the inner margins of com-

pound eyes for Sphinx moths collected in the vicinity of the Tall Grass Prairie Preserve, Manitoba.

ie,

Ceratomia undulosa Harris Sphinx chersis (Hubner)

Sphinx kalmiae J.E. Smith Sphinx luscitiosa Clemens Sphinx drupiferarum J.E. Smith Smerinthus cerisyi Kirby Smerinthus jamaicensis (Drury) Poanes excaecatus (J.E. Smith) Poanes myops (J.E. Smith) Cressonia juglandis (J.E. Smith) Pachysphinx modesta (Harris) Hemaris thysbe (Fabricus) Hemaris diffinis (Boisduval) Darapsa myron (Cramer)

Hyles gallii (Rottenburg)

DUDA OWMAAW® W841

Mean proboscis

Mean distance between outer

Mean distance between inner

length eye margins eye margins (mm + SE) (mm + SE) (mm + SE) 9.14 + 0.76c* 5.05 + 0.05h 2.04 + 0.02d

40.32 + 0.48h 5.89 + 0.03k 2.40 + 0.llef 33.64 + 131g 5.31 + 0.17bi 2.05 + 0.18d 22.56 + O.17f 4.46 + 0.05ef 2.34 + 0.01be 38.40 + 0.97h 5.59 + 0.061 2.34 + 0.02e 2.21 + 0.06a 4.76 + 0.13¢g 1.88 + 0.03c 1.73 + 0.75a 3.85 + 0.03c 1.70 + 0.02b 2.89 + 0.03b 4.63 + 0.05fg 2.09 + 0.03d 1.87 + 0.04a 4.07 + 0.03d 1.83 + 0.06be 2.10 + 0.05a 3.31 + 0.08a 1.37 + 0.53a 1.95 + 0.17a 5.73 + 0.07] 2.53 + 0.02F 12.75 + 0.52d 3.88 + 0.02c 1.88 + 0.08¢ 9.48 + 0.20c 3.58 + 0.06b 1.89 + 0.03c 17.68 + 0.18e 4.28 + 0.09e 2.12 + 0.04d 33.50 + 0.96g 4.88 + 0.03gh 2.44 + 0.02ef p = 0.001 p = 0.001 p = 0.001 14,98 — 37.1 Fi yos = 99-2 F495 = 40.1

‘Number of moths measured.

*Means within each column followed by the same letter are not significantly different (Fishers LSD, p > 0.05).

a proboscis of sufficient length to take nectar from P. praeclara. H. gallii populations fluctuate greatly from year to year in southern Manitoba, often with very few individuals appearing in some years. Therefore, low pollinator populations may be restricting seed produc- tion in some years for orchids in Manitoba.

The Tall Grass Prairie Preserve is surrounded by agricultural lands, a mixture of intensively farmed grains and oilseeds and livestock production. The lar- val host plants for S. drupiferarum can very regionally and include Malus spp., Prunus spp. and lilac, Syringa vulgaris L. (Hodges et al. 1983). These plants are pre- sent in the vicinity but their distribution is patchy. It is unknown whether insecticide usage (Suzan et al. 1994) on adjacent farmlands may restrict the population of larval forms of S. drupiferarum and H. gallii or if weed control may restrict access to host plants. $. drupifer- arum may be a more efficient pollinator than H. gallii, but the number of S. drupiferarum adults frequenting the orchid fields was low in this study. There may be considerable competition between nectar sources given the amount of intensively managed lands adja- cent to the Tall Grass Prairie Bresoive: which would further reduce the time spent by moths pollinating or- chids. Strong light sources from farms surrounding the orchid fields may also ao moths, preventing them from visiting orchids. S. drupiferarum occupies the northernmost extension of its range in Manitoba, which may also contribute to sporadic population oc- currence and lower pollination rates of P. praeclara.

Baker (1961) stated that sphingophilous flowers of- ten have a low frequency of pollinator visitation and compensate by producing numerous seeds. Further research is required to determine if other sphinx moth species are pollinating orchids and if the current level of seedpod production is abnormally low or normal for P. praeclara in Manitoba.

ACKNOWLEDGMENTS

This project was funded by the following agencies: Manitoba Conservation, the University of Winnipeg, Manitoba Orchid Society, Manitoba Sustainable Development Fund and the World Wildlife Fund. The authors would like to acknowledge Jason Greenall, Peggy Westhorpe, Lorne Heska, Liz Reimer and Tracy Loewen in provid- ing assistance and support to this study and Dr. C. Wang and Rachel Boone for reviewing the manuscript. Photographs by R. Westwood.

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Received for publication 30 January 2003; revised and accepted 29 July 2003.

Journal of the Lepidopterists’ Society 58(1), 2004, 21-32

EMPYREUMA SPECIES AND SPECIES LIMITS: EVIDENCE FROM MORPHOLOGY AND MOLECULES (ARCTIIDAE: ARCTIINAE: CTENUCHINI)

SUSAN J. WELLER,' REBECCA B. SIMMONS? AND ANDERS L. CARLSON?

ABSTRACT. | Species limits within Empyreuma are addressed using a morphological study of male and female genitalia and sequence data

from the mitochondrial gene COI. Currently, four species are recognized: E. pugione (L.

), E. affinis Rothschild, E. heros Bates, E. anassa

Forbes. Two entities can be readily distinguished, the Jamaican E. anassa and a ee E. pugione-complex, based on adult morphology. Neither E. affinis nor E. heros can be distinguished by coloration or genitalic differences. Analysis of COI haplotypes suggests that E. affinis is not genetically distinct from E. pugione (<1% sequence divergence); however, the population from the Bahamas, E. heros, is differentiated from other haplotypes with an uncorrected sequence divergence of 5%. We place E. affinis Rothschild, 1912 as a new synonym of E. pugione Hiibner 1818, and recognize three species: E. anassa, E. pugione, and E. heros. This paper includes a revised synonymic checklist of species and a redescription of the genus, with notes on biology, and with illustrations of male genitalia, female genitalia, wing venation, and abdominal

sclerites.

Additional key words: Caribbean fauna, Greater Antilles, mimicry, phylogeography, systematics.

The tiger moth genus Empyreuma Hiibner (Arcti- idae: Arctiinae: Ctenuchini) (Hiibner 1818) is endemic to the Greater Antilles of the Caribbean, and has ex- panded its distribution into Florida (Adam & Goss 1978, Franclemont 1983). Adults are colorful mimics of the wasp Pepsis rubra Drury (Hymenoptera: Pom- pilidae) (Fig. 1A—D), and the host plant for all species reared to date is Nerium oleander (L.) (Apocynaceae). Mating behavior of E. pugione (L.) involves ultrasound signaling between males and females (Coro et al. 1983, Otazo et al. 1987, Perez et al. 1988, Portilla et al. 1987, Wilson 1999). In some of these studies, E. pugione was misidentified as E. affinis Rothschild. Continued con- fusion over the species status of E. pugione, E. affinis, and other members of this genus frustrates attempts to interpret mating experiments among populations ob- tained from different locations in the Caribbean.

Previous taxonomic treatments have been summa- rized in an annotated synonymic checklist by J. Don- ahue (unpublished). Currently, four valid species and two subspecies names are recognized in Empyreuma. These include E. pugione (type species; type locality Virgin co _ E. affinis affinis Rothschild (type local- ity Cuba), E, affinis haitensis Rothschild (type locality Haiti), E. anassa Forbes (type locality Jamaica), and E. heros Bates (type locality Bahamas). Forbes (1917:344) treated E. affinis and E. pugione as separate species when he described E. anassa, but later (Forbes 1930) refers to just two species, one restricted to Jamaica (E. anassa) and one widespread throughout the Greater Antilles. That is, Forbes considered E. pugione and E. affinis conspecific, although he did not formally place E.

‘Dept. of Entomology, 1980 Folwell Ave., University of Min-

nesota, St. Paul, Minnesota 55108, USA.

* Systematic Entomology Laboratory, USDA/ARS/PSI, Smith- sonian Institution, P.O. Box 37012, Natural History Building, E-525,

0168, Washington DC 20013-7012, USA.

*1503 Madison St. NE Apt. 2, Minneapolis, Minnesota 55413, USA.

affinis as a junior synonym of E. pugione. Bates (1934) subsequently described E. heros from the Bahamas, but he did not provide figures or diagnostic features that separate it from previously described species. Thus, the question remains whether either or both of these de- scribed species are junior synonyms of E. pugione.

We report here the results of a morphological sur- vey of genitalia and a molecular characterization of haplotypes of E. pugione, E. affinis, and E. heros. We find that the morphological evidence supports recog- nition of two species, E. anassa and a widespread, ex- ternally variable E. pugione as Forbes (1930) sug- gested. In contrast, haplotype differentiation suggests that the population in the Bahamas is genetically dis- tinct from other populations of E. pugione supporting recognition of E. heros as a third, valid species.

MATERIALS AND METHODS

Morphology. Standard genitalia dissections were done (Winter 2000). Abdomens were softened in warm 10% KOH for 5-15 minutes and then cleaned (scales and viscera removed) in several rinses of 40% ethanol. Abdominal sclerites and genitalia were stained with chlorazole black E (Sigma, St. Louis, MO) dissolved in deionized distilled water (saturated). Specimens were viewed in 40% ethanol. Wings were bleached, neutral- ized in weak acetic acid, rinsed and stained overnight in Eosin Y (1% in distilled water; Fisher Scientific, Pitts- burgh, PA). Permanent slide mounts (Canada balsam, Sigma, St. Louis, MO) were made of abdominal pelts, genitalia and wings (Winter 2000).

Genital preparations of 18 reared individuals (9 males, 9 females) were examined to assess variation within a population (Table 1). These individuals were offspring of pairs of wild caught individuals (W. Con- ner pers. com.). Type specimens of E. affinis affinis Rothschild (BMNH), E. sanguinea Rothschild, E. a. haitensis Rothschild (BMNH; 2 males and 2 female

Fic. 1. Adult males of Empyreuma pugione L. (A), E. heros (B), and E. anassa Forbes (C); Adult female of Pepsis rubra Drury (Hy- menoptera) (D).

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syntypes including #2459), E. a. portoricensis Roth- schild (female syntype 2460) were examined. The E. heros type was not available for loan from the MCZ and its image is not on-line on the type specimen data- base. The species E. anassa is distinct and the uncus sufficiently illustrated by Forbes (1917) to allow confi- dent determination. Additional dissections of speci- mens were made that represented type localities of E. anassa, E. affinis, E. heros, and E. pugione. Camera lu- cida drawings were made of selected specimens. Spec- imen deposition and genitalic preparation numbers are indicated in “Specimens examined” and in Table 1. Wing measurements were taken from the center of the thorax to the wing tip (wing length) and from wing tip to wing tip (wing span).

Terminology for abdominal and genital morphology follows Klots (1970) and Forbes (1939). Collections consulted include: FSMC, Allyn Museum, Florida State University (J.Y. Miller); BMNH, The Natural History Museum, London (M. Scoble); MNHP, Muséum Na- tional d'Histoire Naturelle, Laboratorie d’Entomologie, Paris (J. Minet); NMNH, National Museum of Natural History, Smithsonian Institution, Washington, D.C. (M. Pogue); UMSP, University of Minnesota Saint Paul In- sect Collection (R. Holzenthal).

Gene region and analysis. Thirteen individuals from the Puerto Rican colony were sequenced for the mitochondrial gene COI and these represent E. pu- gione portoricensis Rothschild. Eight individuals were collected from Fort Lauderdale (Florida) and repre- sent E. affinis. Identifications were confirmed at the NMNH by R. Wilson and R. Simmons. These vouch- ers were deposited at the Insect Museum (St. Paul, Minnesota). Museum specimens were used to obtain a set of individuals (9) from the Bahamas, and legs of these were extracted to represent E. heros. Multiple attempts to extract DNA from museum specimens of E. anassa were unsuccessful. We suspect that tradi- tional preparation techniques, drying in paper en- velopes followed by relaxation for spreading, degraded the DNA. Museum specimens were collected over multiple years and a single leg per museum specimen was used. For each individual, the source colony or museum collection, voucher number, and sex are re- ported in Table 1.

DNA extractions were performed using the DNeasy Tissue Kit® (QIAGEN Inc., Santa Clarita, CA) and the Insect extraction protocol (DNeasy Tissue protocol 1997) with 20 ul of Proteinase K (20 mg/ml). Either frozen material (20°C) or legs of museum specimens were used. Museum specimens were ex- tracted on separate days from fresh material to mini- mize contamination with similar DNA. DNA extrac-

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bo (es)

TaBLE 1. Specimens used in mtDNA study. “SJW#” = genitalic preparation of individual, “DNA#” = DNA voucher number for same, E. =

Empyreuma, — = not applicable. Voucher no. Source/label Haplotype GENBANK Voucher Genus species dissection DNA data number number depository E. pugione SJW10542 DNA900 Puerto Rico Colony 1 AF513059 UMSP SJW10552 DNA9QO1 Puerto Rico Colony 2 AF513060 UMSP SJW1056¢ DNA903 Puerto Rico Colony 3 AF513062 UMSP SJW1057¢ DNA904 Puerto Rico Colony 4 AF513063 UMSP SJW1058¢ DNA905 Puerto Rico Colony 5 AF513064 UMSP SJW1059¢ DNA906 Puerto Rico Colony 6 AF513065 UMSP SJW 10622 DNA909 Puerto Rico Colony tf AF513068 UMSP SJW 10632 DNAQ10 Puerto Rico Colony 8 AF513069 UMSP SJW10642 DNAQI1 Puerto Rico Colony 9 AF513070 UMSP / DNAQI18 Puerto Rico Colony ll AF513075 UMSP DNA920 Puerto Rico Colony 3 AF513077 UMSP DNAQ21 Puerto Rico Colony — — UMSP E. affinis 2 DNA902 Florida Colony 12 AF513061 UMSP SJW1060¢ DNA907 Florida Colony 13 AF513066 UMSP SJW10612 DNA908 Florida Colony 14 AF513067 UMSP SJW1065¢ DNAQ12 Florida Colony 15 AF513071 UMSP — DNA9Q13 Florida Colony 10 AF513072 UMSP SJW1066¢ DNAQ14 Florida Colony 16 AF513073 UMSP SJW 10676 DNAQ17 Florida Colony 17 AF513074 UMSP SJW1069¢ DNAQI19 Florida Colony 18 AF513076 UMSP DNA922 Florida Colony 19 AF513078 UMSP E. heros SJW1081¢ DNAS043 Bahamas: Long Island 21 AF513083 FSMC SJW1082¢ DNAS038 Bahamas: Crooked Island 20 AF513080 FSMC SJW10832 DNASO41 Bahamas: Crooked Island 21 AF513081 FSMC DNAS042 Bahamas 22 AF513082 FSMC Nyridela sp. — DNA069 Las Alturas, Costa Rica _ AF513079 UMSP Scena potentia — DNAOOS Las Alturas, Costa Rica = AF277448 UMSP

tion control blanks were maintained for each museum

(94°C, 1 min, 45°C, 1 min, 72°C, 10 min), 4°C for a

extraction set. These blanks were checked for volatile DNA contamination by including them in the PCR amplifications. All extraction blanks were negative (did not contain DNA) when used as template for PCR.

The entire COI gene was amplified using PCR (Saiki et al. 1988) for lab colony individuals (Table 1). The COI primers amplify nearly 1500 bp of COI. To amplify COI, five primers (two external, three inter- nal) were used. The external primers were K698 (5’- TAC AAT TTA TCG CCT AAA CTT CAG CC-3’) and PAT2K837 (5’-TCC ATT ACA TAT AAT CTG CCA TAT TAG-3’) that have 5’ ends located at positions 1436 and 3037, respectively, on the Drosophila mt genome (Clary & Wolstenholme 1985). Three internal primers were also used: C1-J-1751 (alias RON), C1-N- 2191 (alias NANCY), and REVNANCY (5’-GAA GTT TAT ATT TTA ATT TTA CCG GG-3’; position at 5’: 2190) (Simon et al. 1994).

Based on initial results of haplotype variation, only the more variable portion of COI, a 550 bp piece (revNancy-Pat2K837), was amplified and sequenced for specimens from the Bahamas (E. heros). For all re- actions, a hot start (95°C dwell, 1 min) prior to addi- tion of TAQ was used. Cycling parameters were: 29 cy- cles (94°C, 1 min, 45°C, 1 min, 72°C, 1 min), 1 cycle

minimum of 4 minutes. PCR products were cleaned for automated sequencing with a Qiaquick PCR purifi- cation kit® (QIAGEN Inc., Santa Clarita, CA) accord- ing to protocol. Sequencing reactions were performed using Bigdye terminator kit (PE Biosystems) using 10 um of primer and 1-6 ul of clean PCR product. Se- quencing reactions were performed using a BigDye Terminator Cycle Sequencing Ready Reaction Se- quencing Kit® (PE Applied Biosystems, Foster City, CA). We performed half reactions and used 2 UL of 10 uM sequencing primer, 1-6 WL of clean PCR product, and $-13 wL ddH,O (final volume: 20 uL). Recom- mended sequencing cycling parameters were used. Each sample was cleaned using Sephadex columns (Centri-Sep protocol; Princeton Separations, Inc., Adelphia, NJ). Samples were then resuspended in 20 uL of Template Suppression Reagent (TSR)® (PE Applied Biosystems, Foster City, CA). An ABI 310 sys- tem was used to visualize and record the sequence. Typically, sequences up to 750 bp were obtained with the long capillary for COT.

Data were imported into Sequencher 3.1.1® (Gene Codes Corp., Ann Arbor MI). Sequences for each in- dividual were aligned to produce a consensus se- quence and the sequence translated and checked for

stop codons. Individuals were then aligned by con- served motifs and adjusted by eye when necessary.

Phylogenetic analysis. As the relationship of

Empyreuma to other ctenuchines and euchromiines is

unknown, we established potential outgroups by per-

forming an initial analysis with one sequence of Empyreuma and all available ctenuchine and eu- chromiine species (Simmons & Weller 2001). We then selected species of two genera, Nyridela sp. and Scena potentia (Druce), to root the analysis of Empyreuma haplotypes. These taxa are given in Table 1 with their GENBANK accession numbers. Unique haplotypes were identified for the Empyreuma specimens. These were analyzed using heuristic searches and maximum parsimony (PAUP*) (Swofford 2000). All positions were equally weighted, and 10 random additions were performed to search for tree islands (Maddison 1991). The parsimony results were then used to generate likelihood parameters for a maximum likelihood analy- sis (ML) using the following menu options: Trees: Tree scores: Likelihood. The following likelihood parame- ters were used: HKY-85 model, transition-transversion ratio of two, and empirical nucleotide frequencies.

RESULTS AND DISCUSSION

How many species? Morphology clearly supports the recognition of two entities, E. anassa, E. pugione- complex, whereas molecular results supports three species, E. anassa, E. pugione, and E. heros. The male genitalia (Figs. 2-3) and female genitalia (Fig. 4) of E. anassa and the E. pugione species-complex are distinct. However, we could not identify consistent, adult mor- phological features to diagnose E. heros compared to E. pugione (compare Fig. 2B, C). Both coloration and armature of the male vesica lack fixed differences.

Our molecular results differentiate between E. heros (the Bahamas), and other E. pugione popula- tions. The mtDNA sequences of Empyreuma were typical for COI in insects (Simmons & Weller 2001), with an A/T bias, especially pronounced in the third codon position (A = 43%, C = 12%, G = 2%, T = 43%). We obtained approximately 1474 bp of COI sequence for E. pugione and E. affinis, and 450 bp for E. heros, from approximately 2190 to 3037 (revNancy-Pat2K837). Of 1474 bp, 54 bp were informative (4%); the majority of this variation was third positions (first: 10/54, sec- ond: 5/54, third: 39/54). We obtained 19 distinct hap- lotypes for the combined sample of E. pugione and E. affinis; three were obtained for E. heros. The uncor- rected p-distance between E. heros to the other haplo- types was 5%. There are 17 unique substitutions for E. heros (Table 2), and all are third position transitions.

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The MP analysis of the COI data resulted in over 139,000 trees (length = 302, consistency index = 0.66, retention index = 0.69: trees not shown). The ML analysis (Fig. 5) ine a —In L = 3571.17. Individuals from the Puerto Rico population (E. pugione portori- censis) and from the Florida population (E. affinis) do not segregate into two, reciprocally monophyletic taxa (Fig. 5). In contrast, the haplotypes from the Bahamas are recovered as a separate clade in all observed COI topologies. Cryptic species in leaf-mining flies and other insects have been identified by COI and other molecular markers (e.g., Frolich et al. 1999, Scheffer 2000, Scheffer & Lewis 2001). The genetic divergence between E. affinis and E. pugione (<1%) is slightly higher than divergences among races of Heliconius er- ato (0.5%; Brewer 1994a, b) or agromyzid flies (0.6%; Scheffer & Weigmann 2000); however, there is no clustering pattern to the pugione-affinis haplotypes (Fig. 5). Similarly, a study of Western spruce budworm species’ limits (Sperling & Hickey 1994) suggested that the designation of Choristoneura biennis Free., C. orae Free., and C. occidentalis Free. could not be sup- ported because of low sequence divergence (<1%), and their haplotypes were placed in the same clade.

Both morphological and molecular results support placing E. affinis as a junior synonym of E. pugione. Molecular results support maintaining the species’ sta- tus of the Bahaman population, E. heros. Additional sampling and molecular study is warranted to confirm these results; however, these initial findings support treating this population as a unique, endemic lineage that should be considered when forming conservation strategies for the Bahamas. Additionally, other islands in the Caribbean may harbor cryptic genetic diversity and greater sampling is needed.

Phylogenetic placement. The systematic place- ment of Empyreuma is unclear. Although a prelimi- nary study of Ctenuchini and Euchromiini identified Scena and Nyridela as potential sister genera (Sim- mons & Weller 2001), that study was focused on as- sessing the monophyly of tribes, not identifying the nearest sister genus or genera to Empyreuma. Our COI results suggest that these genera, Scena and Nyridela, are not sister to Empyreuma. Rooting trees with Scena and Nyridela places the root at the mid- point of the longest branch: the branch connecting the E. heros clade to the E. pugione haplotypes (Fig. 5). That is, using Scena and Nyridela as outgroups was no better than arbitrarily selecting midpoint rooting for our analysis. Morphological data does not support a close relationship either. Males of Scena potentia Druce have a bifid uncus apically, but it is stalked at the base and probably not homologous with the bifid

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wo 1

Fic. 2. Male genitalia of: E. pugione culture, SJW1066 (A), E. pugione Dom. Rep., SJW1005 (B), E. heros, SJW1001, (C), E. anassa, Ja- maica, S)W999 (D). A = anal tube, J = juxta, M5a = muscle attachment process, S = sacculus, SP = spinose pad, T = transtilla, U = bifid uncus, V = vinculum.

Fic. 3.

condition of Empyreuma (not figured). Male genitalia of Nyridela chalciope Hiibner have a single uncus and differ in other genitalic details (not figured).

To place Empyreuma in the ctenuchine-euchromiine lineage, we surveyed the morphology of other ctenu- chines and euchromiines (Weller et al. 2000). Females of Euchromia lethe Fabricius possess main and acces- sory bursae, but the corpus bursa has the anterior third lined with spiny setae, not organized into distinct strips (SJW928 NMNH). Females of Napata albiplaga Walker, N. splendida Herrich-Schiiffer, and Cyano- pepla bella Guérin also possess a main and accessory bursa with spination similar to E. lethe, but differ in spine placement and numbers. All of these females and Empyreuma females share the flattened ovipositor lobes and highly wrinkled, melanized membrane surrounding the ovipore. Males of these species do not share obvious similarities with Empyreuma, but this is merely a phenetic assessment. A cladistic analysis with more exhaustive sampling of ctenuchines and eu- chromiines, using both morphological and molecular data, will be needed to establish generic relationships

TABLE 2. Drosophila COI gene (Clary & Wolstenholme 1985).

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C

Aedeagus of E. pugione (SJW1005). C = cornuti, V = vesica.

and tribal limits (Simmons & Weller 2001, Jacobson & Weller 2002).

Taxonomy. Below is a redescription of the genus Empyreuma. We provide illustrations of the habitus, wing venation, and male and female genitalia of the two morphologically distinct species, E. pugione and E. anassa. An abbreviated species diagnosis and de- scription is provided for E. heros.

Empyreuma Hibner, 1818

Chrysaor Hiibner, 1808:4. Type: Chrysaor erythropterus Hiibner1808:4, designated by monotypy. Work rejected for nomenclatural purposes by ICZN, 1966, Opinion 789. Chrysaor was placed on Official In- dex of Rejected and Invalid Generic Names in Zoology (Wat- son et al., 1980:61).

Empyreuma Hiibner 1818:12. Type species: Sphinx pugione Linnaeus, 1767:807, by subse- quent designation by Hampson, 1898:422.

Diagnosis. The coloration of species of this genus (Fig. 1A-C) distinguishes them from other ctenu- chine/euchromiine genera. The blue-black body and black-fringed, scarlet wings is unique among the

Sites of unique substitutions for E. heros compared to E. pugione haplotypes. Numbering corresponds to the position on the

Site numbers: 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 9 7 7 vii 0 x 8 8 8 8 9 9 ) 9 0 0 0) 1 4 6 6 8 9 3 dh ri 9 6 8 9 9 0 3 6 5 6 1 4 2 2) 0) 2 8 3 5 6 ® § 4 7 1 4 E. heros A ily Cc C C A G Cc A Cc A Cc G C G T Cc E. pugione C G at A T G ‘ar A (G ae ‘a a aw T oT: G ‘a

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Fic. 4, Female genitalia of: E. pugione Puerto Rico culture, S]W1054 (A), E. pugione Puerto Rico culture SJW1068 (B), ductus and corpus bursae of E. anassa, Jamaica, SJW997 (C), details of lamellae postvaginalis (LPV) of E. pugione SJW1068 (D) and LPV E. anassa Jamaica, SJW997 (E). AB = appendix bursa, ADB = accessory ductus bursa, AP = anterior apophyses, C = corona of cornuti, CB = corpus bursa, DB = ductus bursa, DS = ductus seminalis, O = ostium, PA = papillae anales, PP = posterior apophyses, S = signa, VIII = Sth sternite, V = vesica.

mimetic species (RBS pers. obs.). The bifid uncus of spinose signae of the females (Fig. 4S) are also distinc- the males (Fig. 2) is diagnostic for Empyreuma, and ap- tive and possibly derived only in this genus. pears to be unique within the Euchromiini-Ctenuchini Adult habitus (Fig. 1A-C). Wings opaque with brown, black or

clade (sensu Jacobson & Weller 2002). The paired, blue-black scales on upper surface of forewing. Relatively large

affinis 913 pugione 903 pugione 920 pugione 918 pugione 906 > pugione 911 pugione 909 ugione 904 ‘prigions 905 pugione 910 pugione 901 affinis 914 pugione 900 affinis 919 affinis 917 affinis 902 affinis 912 affinis 922 affinis 908 affinis 907 heros 38 heros 41 heros 42 Nyridela

Scena

Fic. 5. Maximum likelihood tree with Neridela and Scena as outgroups. Taxon name includes specimen DNA voucher number (see Table 1).

moths, average male wingspan 49 mm and average wing length 21.8 mm (n = 82). In females, average wingspan 50 mm and average wing length 20.5 mm (n = 90). ). Unipe ctinate antenna with a black shaft for nearly entire length contrasting with orange apex. Ground color of head, thorax, and abdomen Blaale blue Miele or brownish depend- ing on specimen.

Head and Thorax. Antennae unipectinate; ocelli present with a melanized outer ring. Proboscis longer than head. Prothorax lacks parapatagia or a dorsal gland; ephiphysis short. Both meso- and metathoracic legs possess simple claws (not bifid), and tibial spurs with smooth (not serrated) apices.

Wings (Fig. 6). Forewing venation: Sc slightly sinuous, ex- tending nearly 9/10 of costa. R, and R, arise from discal cell, and R,. ste ked: with R. arising closer to cell than R,-R, branching. M, arises from cell and separate from R, .. The cross ‘vein between M, and M, typically thins or even has a break anterior to M.,. M, arises ines M. with a short spur of M, extending into cell. C uA, andl CuA, wide ly separated and extend to wing edge. Hindwing venation: Sc + R absent. RS and M, connate. Discal cell cross-vein strongly devel- oped and V-shaped. M, weakly developed, and extends to wing mar- gin. M, strongly dev eloped, Tee from apex of discal cell. CuA, and C uA, stalked. and Anal vein ( A) just touching wing margin. Fe “ells with two frenular bristles.

Abdomen. In both sexes, second sternite with long, straight apodemes (Fig. 7), and 2nd and 3rd sternites and tergites ‘Jack mod- ifications for wasp mimicry (Weller et al. 2000). ). Male. lacking andro- conia (no abdominal scent pouch or coremata). Sth sternite S weakly sclerotized (Fig. 7).

Genitalia. iiales (Figs. 2, 3): Tegumen relatively short, arising nearly perpendicular to the vineullneiny uncus bifid, long processes

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with rounded apices (Fig. 2A—C), or bifid processes short with pointed apices (Fig. 2D); narrow, sinuous vinculum ventrally pro- duced as rounded shallow saccus. Valve with strongly sclerotized base and costa; attachment point of M5 (valve extensor muscle; Forbes 1939, Tikhomiroy 1979) marks edge of costa and sacculus (Fig. 2, M5a); sacculus extending apically as a membranous lobe (distinct from costa) bearing a large, spinose pad on internal surface (Fig. 2A-C, SP; E. pugione-complex), or spinose pad reduced to flat- tened, irregular, rugose area fused to costa (Fig. 2D, SP; E. anassa); membranous ventral edge of sacculus with few setae and sclerotized base coincident but not fused to juxta. Juxta protruding posterio- rally in a bell-shaped projection with wishbone-shaped thickening of edges (Fig. 2A-D); Anellar region with small anellar sclerites

fused to venter of aedeagus, pair of slender, crescent-shaped patches lie dorsad of aedeagus (Fig. 2, T); anal tube (Fig. 2A) with pair of irregularly shaped selenite »s or anal tube lightly sclerotized. Aedea- gus (Fig. 3) relatively large, compared to genital capsule; endophal- lus with a sclerotized tube possessing a flattened apex ringed with short spines (Fig. 3); varying number of teeth-like spines, and num- ber not corresponding with species-limits.

Females (Fig. 4A—D): Papillae anales (PA) lightly sclerotized, laterally flattene d; membrane surrounding ovipore highly folded with melanized striations; posterior apophyses (PP) long and narrow; dorsal pheromone glands as paired narrow tubes with rounded or crescent-shaped thickenings at midpoint and terminus—as long or slightly longer than posterior apophyses (not shown). 8th tergum and sternum fuse at right angles with very short anterior apophyses (Fig. 4B, AP); 8th tergum aoauitils d, broad with anterior e dge highly concave; 8th stemum weakly sclerotized and lacking ornamentation (Fig. 4A-B, detail 4D; E. pugione complex), or atl a distinct, rounded lamella postvaginalis in E. anassa (detail, Fig. 4E); lamella antevaginalis absent; ostium bursa marked by a membranous ventral

edge of ductus bursa. Ductus bursae (DB) short, membranous and same width as ostium bursa. Corpus bursae (CB) with two oblong, highly spinose signae located opposite one another ventrally atl dorsally, and nied to an accessory bursa by a twisting duct. Ductus seminalis (DS) arising from accessory ductus bursa (ADB). Acces- sory bursa (AB) comprised of thinner membrane, lacking orna- mentation.

Notes on Biology. Larvae of these species have been reared on Nerium oleander (Apocynaceae) based on museum labels. The plant genus Neriwm has only three species, and is native from the Mediterranean to Japan. The exotic species, N. oleander has been natu- ralized widely in North America (Correll & Johnston 1979) and it contains cardiac glycosides. The native hosts of Empyreuma have not been recorded, but po- tential New World apocyanaceous host genera include Thevetia L., Plumeria L., Mandevilla Lindl., and Tabernaemontana L..

Empyreuma anassa has been collected from nearly sea level to 918 m elevation. Similarly, E. pugione has been collected up to 733 m. Adults have been col- lected in April, May, July, September, and December. Historical label data for E. heros lacks elevation, but flight times are similar occurring in October, Decem- ber, and March.

Discussion. Empyreuma anassa is easily separated from the E. pugione complex, based on color and gen- ital differences. Our examination of the male genitalia suggests that intraspecific variation exists in the shape of the uncus, presence and development of a spinose

VOLUME 58, NUMBER 1

Fic. 6. Wing venation of forewing (below) and hindwing (above). A = anal vein, CuA, = cubital vein, f = fold, R = radial vein, Rs = radial

sector, Sc = subcosta, 1-5 = vein number.

patch on the sacculus, and presence of a small tooth on the vesica for specimens identified as either E. pugione or E. affinis (variation not shown). Specimens of the subspecies described by Rothschild (E. a. haitensis, E. a. portoricensis) fall within the ee of morphological variation observed in the reared cultures from Florida and Puerto Rico. Both are placed as new junior syn- onyms of E. pugione. These subspecies may not be de- fensible: however, more complete survey of moths, their larvae, and their DNA across the Greater Antilles is needed before subspecific status can be discarded de- finitively, work that is in progress (J. Rawlins in prep).

Empyreuma pugione (Linnaeus, 1767)

Empyreuma pugione (Linnaeus, 1767).

Sphinx pugione Linnaeus, 1767. Syst. Nat. (Ed. 12) 1(2):807. Type Locality: insula S. Thomae [St. Thomas Island, Virgin Islands]

Sphinx lichas Cramer, 1775. Papillons Exot. 1:70; pl. 45, fig. B. Type Locality: St. Thomas, Virgin Islands [not Zygaena hie has Fabricius, 1781, from Arabia, in the Zygaenidae].

Sphinx sanguinosa Martyn, 1797. Psyche: pl. 26, figs. 18, 19 (un- available?).

Chrysaor erythropterus Hiibner, 1808. Erste Zutrage Samm. Exot. Schmett.: 4. Unavailable: sole species included in Chrysaor, in a work rejected for nomenclatural purposes.

Empyreuma lichas: Hampson, 1898, not Fabricius, 1781. Cat. Lep. Phal. Brit. Mus. 1:423; fig. 223. (Misidentification).

elie mucro Zerny, 1912 (25 July). In Wagner, ed., Lepid.

Catalogus 7:122 [unnece ssary replacement name for Zy- gaena elias sensu Hampson, 1898, not Fabricius, 1781].

Empyreuma sanguinea Rothschild, 1912 (21 Dec.). Novit. Zool. 19:155. [unnecessary replacement name for Zygaena lichas sensu Hampson, 1898, not Fabricius, 1781].

Hampson, 1914. Cat. Lep. Phal. Brit. Mus. Supp. 1:267 [as valid name for taxon Hampson, 1898:423 had misidentified as E. lichas].

Forbes, 1917. Bull. Amer. Mus. Nat. Hist. 37:344 [as syn- onym of E. pugione].

Empyreuma sanguinea portoricensis Rothschild, 1912 (21 Dec.). Novit. Zool. 19:155. Type Locality: Puerto Rico.

Hampson, 1914. Cat. Lep. Phal. Brit. Mus. Supp. 1:267 [as synonym of E. sanguinea).

Empyreuma affinis Rothschild, 1912. Novit. Zool. 19:155. Type Locality: Cuba; new synonym.

Forbes, 1917. Bull. Amer. Mus. Nat. Hist. 37:344. Hampson, 1914. Cat. Lep. Phal. Brit. Mus. Supp. 1:267: pl. 13, fig. 31.

Empyreuma affinis affinis Rothschild, 1912. Novit. Zool. 19:155. Type Locality: Cuba; new synonym.

Empyreuma affinis haitensis Rothschild, 1912. Novit. Zool. 19:156. Type Locality: Haiti; revised synonym [of E. pu- gione (L.)].

Forbes, 1917. Bull. Amer. Mus. Nat. Hist. 37:344 [as valid “race” of E. affinis; misspelled as “haytiensis”].

Hampson, 1914. Cat. Lep. Phal. Brit. Mus. Supp. 1:267 [as synonym of E. affinis].

Empyreuma haytiensis Forbes, 1917. Bull. Amer. Mus. Nat. Hist. 37:339, 344. Misspelling.

Diagnosis. The wing length in males ranges from 19-27 mm (A = 21.8: STD = 2.5: n = 24), and is 17-27 mm in females (A = 19.9: STD = 2.4: n = 35). The forewing varies from primarily brownish red with

30

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Fic. 7. Male abdominal sternites of E. pugione (SJW1005): sec- ond (II) above, eighth (VII) below.

costal red streak (Dominican Republic, Haiti) to brownish red with orange costal streak (Cuba), and has a thin black border on the outer margin (Fig. 1A, B). The forewing lacks scarlet coloration on the un- derside. The black border varies in width between in- dividuals. White markings are present on the thorax, the patagia, the tegulae, and the wing bases. These white markings are more strongly developed in speci- mens from Bahamas. Medial, dorsal white patches are present on Al and A2. There is a small pair of white spots on A2 in pleural region. Conspicuous, paired white patches are present on the dorsum of A3 and A4. Smaller than those occurring on A3 and A4, white patches occur on the anterior edge of A5—A7. Pleural spots are present on A3-5 and A7-8. These white patches are missing or reduced in some specimens and patch size varies throughout the range. The tegu- men is relatively short, arising nearly perpendicular to the vinculum. The uncus is bifid, possessing long processes with rounded apices (Fig. 2A—C). The sac-

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

culus extends apically as a membranous lobe (distinct from costa), and bears a large, spinose pad on internal surface (Fig. 2A—C, SP), The anal tube has a pair of ir- regularly shaped sclerites (Fig. 2A). The Sth sternum is weakly sclerotized, and lacks ornamentation (Fig. 4A—B, detail 4D),

Specimens examined: (see Table 1 for additional genitalic dis- sections). CUBA: Baracoa, Coll. Wm Schaus, March (5¢; NMNH). Cayanas, EA. Schwarz (1é; NMNH). Holguin, no other data (1¢, 2°; BMNH):; BM2663 (1d; BMNH); ex Ges Franek 1911 (19; BMNH); H.S. Parish (12; BMNH). Hayana, Baker (26; NMNH). Loma del Gato, Sierra Maestra Cuba, 2500’, July-August [19]29, HF Clement (2¢, 1F; BMNH); same locality: BM2465 (1d, BMNH). Santiago, F. Clement (16; NMNH). Parish affinis syn- types (22; BMNH); 58-126 (12; BMNH); 100-20 (1d, 12: BMNH);: 61-21(1¢; BMNH); affinis syntype 2455 (1d, BMNH). BRITISH VIRGIN ISLANDS: Guiana Island, 1-14 July 1984, SE & PM Miller (1¢: gen. prep. SJW1004; FSMC). Tortola, March [19]66 JAC. Greenwood (12; BMNH). DOMICAN REPUBLIC: La Vega Prov., Hotel Montana, ca. 520 m, 10 km NE Jarabacoa, 28 May 1973, Don & Mignon Davis (1¢: gen prep SJW1005, NMNH). CAYMAN ISLANDS: Georgetown, T.M. Savage English 1911-89 (1¢; BMNH); no other data (12, BMNH). GUADELOUPE: Aer. Du Raizet, ix 81 R. Giscointeties (129; BMNH). HAITI: Petionville, 1300’ 26 Dec. 1954 A. Zerkowitz (1°, BMNH). Port au Prince, 24 X [19]09 Georges Lion, Museum Paris (23; MNHP); same locality, 26 X [19]09 Georges Lion, Museum Paris (1¢; MNHP); same locality, 28 X [19]09 Georges Lion, Museum Paris (1d; MNHP); F. Odile Joseph (1¢, 12, 16: gen prep 2462; BMNH); No. 54-28, Nov. 26, 1928, A. Audant coll. (1¢: gen prep SJW1003; NMNH). Le Perchoir, 3000’ 21 Dec. 1954 A. Zerkowitz (19: BMNH). same locality, 23 Dec. 1954 A. Zerkowitz (1¢; BMNH); same locality, 26 Dec. 1954 A. Zerkowitz (23, 32; BMNH); G. Babault (12; BMNH). San Domingo, syntype F affinis haitiensis (12; BMNH); same locality, syntype ¢ affinis haitiensis (1d; BMNH); Haiti, syntype ¢ (BMNH); Haiti 98, BM2459, syntype °? (BMNH). HONDURAS: (1¢; BMNH), Haiti (26; BMNH). ST. DOMINGO: Tweede 55-1, BM2426 (1d, BMNH). NO DATA. Ex Musaeo Arch. Guenee (1¢é; MNHP); Brazil Ex. Coll. Smith 1894-5 (19; BMNH); America (12; BMNH); Savanah Zyganea lichas (12; BMNH); Zy- ganea pugione Haynes S.P. Oss-Terra (12; BMNH). No label (2¢, 19; BMNH), Empyreuma pugione L. (1°, BMNH); pugione L. (19; BMNH); (2c: NMNH); “Type of sanguinea from S. Domingo?” (1; BMNH); Coll. Bryk Mus. (3¢; NMNH); ornh 2 on 11/69 (1¢; BMNH).

Empyreuma heros Bates, 1934

Empyreuma heros Bates, 1934. Occ. Pap. Boston Soc. Nat. Hist. 8:137. Type Locality: Bahamas (Mariguana Island).

Diagnosis. This species can be diagnosed based on its collection locality and its COI sequence. Unique substitutions are given in Table 2.

Description. Same as E. pugione.

Specimens examined. BAHAMAS: Crooked I, vic Pitts Town, 25 ix 1986 M. Simon & L. Miller, Sta. 1986-UV (1¢: gen prep SJW1001, 12: gen prep SJW1083; FSMC); 1 mi E. Colonel Hill, UV, 18 ix 1988, LD Miller & MJ Simon, Sta. 1988-45, Acc. 1988-18, (1d: gen prep SJW1082; FSMC). Grand Turk Is[land], Xii-1. 1965-66 (12: BMNH). G[reat] Inagua I[sland], Horse Pond ca 1.5 km E of Matthew Town, 28 ix 1986, M. Simon & L. Miller, Sta. 1986-19 (1d gen prep, SJW1002; FSMC). L. Abaro, Mar 1902 (12; BMNH). Long Island, Stella Maris, UV, 26 ix 1988, L.D. Miller & M.J. Si- mon Sta. 1988-63 (2d: gen prep SJW1000, SJW1081; FSMC). Nas- sau I[sland], 19 ii [19]02, J.L. Bohnhote (24, 22; BMNH); J.L.

VOLUME 58, NUMBER 1

Bohnhote (12: BMNH); I. 22 X [18]98 (12: BMNH); G. Carter 1903-6 (12; BMNH); 30 Dec. 1956, A. Zerkowitz (1¢é; BMNH). New Providence Is., FE. Taylor (1¢, 12; BMNH).

Empyreuma anassa Forbes, 1917

Empyreuma anassa Forbes, 1917. Bull. Amer. Mus. Nat. Hist. 37:343; fig. 5. Type Locality: near Troy, Jamaica. Empyreuma pugione: Hampson, 1898 (not Linnaeus, 1767). Cat. Lepid. Phal. Brit. Mus. 1:423. Misidentification.

Diagnosis. E. anassa appears slightly larger than E. pugione, because the body is more robust. The male wing length ranges from 16-26 mm (A = 21.1; STD = Us x n= 5G). Fommale wing length ranges from 16-25 mm (A = 21; STD = 2.0; n = 44). Like E. pugione, the upperside of the forewing is opaque with metallic blue-black or brown scales. In E. anassa, however, the underside is scarlet with a large black border on the outer margin, and the scarlet coloration is visible dor- sally (Fig. 1C). Unlike E. pugione, white scales are lacking on the thorax and legs in E. anassa. The ab- domen has two white crescent bands on A4 that ex- tend to the spiracles but do not meet dorsally. There are smaller, paired bands on A5—A7. Male genitalia also differ between the two species. The bifid uncus of E. anassa is short with pointed apices (Fig. 2D) com- pared to E. pugione (Fig. 2A-C). The spinose pad on the sacculus is reduced to a flattened, irregular, rugose area, which is fused to the costa (Fig. 2D, SP). The anal tube is lightly sclerotized in E. anassa, but not E. pugione. In females, the 8th sternite has a distinct, rounded lamella postvaginalis (Fig. 4E) compared to the trapezoidal one of E. pugione (Fig. 4D).

Species distribution. The species is found in Ja- maica. Three specimens had erroneous label data: British Guiana, Cuba and Costa Rica. The “Mon- eague’ locality is a famous collecting locality in St. Ann Parish Jamaica (Brown & Heineman, 1972), not Costa Rica. The other two specimens lack additional infor- mation (collector or expedition) that would allow de- finitive assignment of locality. These are treated as mislabeled because no other specimens of E. anassa have been taken outside of Jamaica.

Specimens examined (also Table 1). BRITISH GUIANA: Paruima, 26. 10. 38 (12; BMNH). COSTA RICA [Jamaica St. Ann]: Moneague, I iv 1926 1000 ft. F. W. Jackson (12; BMNH). CUBA: Holguin, (1¢; BMNH). JAMAICA: Axe Town, Bred 7.8.[18]92 (Taylor) (1°; BMNH). Bath, USNM Acc 40269 (12; NMNH). Ja- maica: Batton Falmouth, AGM Ja—Feb. 1967 (1d; BMNH). Clar. Par., Portland Ridge, nr Jackson Bay Cave, 40 ft 4 May 1973, Don & Mignon Davis (54, 1d: gen prep S]W999: NMNH); Mason River Station, 4 mi NW Kellits, 2200 ft 16-19 April 73, Don & Mignon Davis (1d, 1d: gen prep SJW1084; NMNH); 2 Km S. Rocky Pt, nr Jackson Bay Cave, Dec 10, 1975 5m, Don & Mignon Davis (2¢; NMNH). Constant Springs, e. xii 1904 Wlsm. (1°; BMNH). Cuna Cuna Pass, Capt. U. Robinson Collector 10 July [19]05 (1M; NMNH). Cornwall County, Coll. Miss M.S. Savarian (19; NMNH). Kingston, 10-1-[18]94 at light (Taylor) (1°; BMNH); at

3]

Ean bred 2.8.[18]92 ( (Taylor) (1¢; BMNH); Taylor) (1d, 12; BMNH); bred 7 8.[18]92 (Taylor) (1d; BMNH); bred pupated 7 7.7.[18]92 emerged 20.7.[18]92 no. 10 E. pugione (Taylor) (1 » BMNH); ); WJ Kaye B. M. 1930-185 (12 BMNH): at light 29-7. -[18}92 (1 > BMNH); 15-IV-73 (1¢; NMNH)- W.R. Maxson Coll. V-29-04 (1d: NMNH). Mandeville, 2000 m July

1923 A. Hall (1d; BMNH); L. J. Bertram (1d; aes Mile Gully, 16 ii 1921 No. 311 (12; BMNH). Moneague, beg. Feb. 1905 Wlsm. (1°; BMNH). Montego Bay, xiii 1923 Major Gillett BM 1924-174 (22; BMNH): 26 Feb. 1911 Miss Fountaine (2d, 12; BMNH). Moore Town, (1¢: BMNH). North Coast, s. level 20 iii 1961 (12: BMNH): s. level 29 i iii 1961 (1d; BMNH): s. level 8 vi 1961 (2¢, 12: BMNH); Dark hours s. level 20 i iii 1961 (26; BMNH): Dark hours s. level 29 iii 1961 (1d; BMNH). Ocho Rios, Dec. 10, 1957 A. Zerkowitz (1d; BMNH): Dec. 28, 1957 A. Zerkowitz (12: BMNH): Jan. 3, 1957 A. Zerkowitz (1¢, 12; BMNH); Jan. 5, 1957 A eae (1d; BMNH); Jan. 6, 1957 A. Zerkowitz (1d, 1; BMNH). Portland Parish, (1¢; BMNH); Hardwar Gap, “Green Hills”, July 24-25, 1962, Farr. O & R. Flint (5d, 12; NMNH): 4 mi. S. Hartford, 850 ft. 26-27 April 1973, Don & Mignon Davis (1°; NMNH). Runaway Bay, end Feb. 1905 Wlsm (1°; BMNH); 5-III-1970, JFG Clarke (12: NMNH). St. Ann Par., Martin (1d; BMNH); Rose Hill, hae Bay 900 ft, 29 April-2 May 1973, Don & Mignon Davis (13¢, NMNH); nr Runaway Bay 50 ft, 1-2 May 1973 , Don & iteeer

Davis (6¢, 12; NMNH). St. Cath. Par., Mt. Diablo Hollymount, 2754 ft 21-24 April 73, Don & Mignon Davis (5d, 12, 1c: gen prep SJW998; NMNH). Westmor. Par., Negril, Dec. 12, 1975, Don & Mignon Davis (2¢; NMNH). BRITISH WEST INDIES: N Coast, S level 20 iii 1961 (2¢, 12; BMNH); same locality: 29 iii 1961 (1¢, 12: BMNH); same locality: 8 vi 1961 (2d, 12, 1d: gen prep SJW996: BMNH). ST. THOMAS: L. Litus, pugione (19; BMNH); Jamaica collection Wm Schaus (5d, 39; NMNH): Female vial #083 Empyreuma anassa R.E. Dietz 1968 (12, NMNH); Jamaica WI, A. Arinoff Donor, July 24, 1933 (1d; NMNH); Cockrell (12: NMNH);: St. Thomas Col. Neumégen, Coll. Brklyn Mus. (1¢; NMNH): Empyreuma pugione 4. 8. 25 Ex. Coll. Griffiths (19; BMNH); LS acti pugione Jamarque (16; BMNH); (60, 8°; BMNH); Tay- lor (63, 49; BMNH); Jamaica Yates (1¢; BMNH); 1960 pres. by George Newman Brit. Mus. 1961-52 (19; BMNH); 40-4-3-104 (1d: BMNH); 45-110 (1d, 12; BMNH); 46-121 (1d; BMNH): 76-71 (1¢: BMNH): E Coll. Hanson (1¢; BMNH); ex Stevens (12; BMNH);: F. W. Jackson 1913-208 (1d; BMNH); F. W. Jackson 1920-341 (1c: BMNH); Ianson (12; BMNH); JMS and J Yates BM 1926-393 (1d; BMNH); ex. Percy I. Lathy 1902 (16; BMNH); (1c: BMNH); P.H. 6-91 Taylor (23, 12; BMNH); R. Stanway Paris 23-ii-1919 (19; BMNH): same data 2-ii-1919 (2¢, 12; BMNH);: same data 16-2-1901 (12; BMNH); West Indies Malrun B. M. 1933-489 (1d, 12: BMNH): Malvun (16; BMNH); British Guiana (16; BMNH); San Domingo 120 (1d; BMNH). NO DATA: Bred 13. 7. [18]92 No. 10 E. pugione (1d; MINTS Bred 25. 7. [18]92 No. 10E. Buco 12: BMNH): 44-11 (1d; BMNH); E pugione no. 10 Bred 16. 7. [18]92 (12: BMNH): Raraia or Jamaica ? (12; BMNH) S. ‘Aaa (12; BMNH); label unreadable (1d; BMNH).

light (Taylor) (1 bred 6.8.[18]92 pil

ACKNOWLEDGMENTS

We thank Julian Donahue for all his generous assistance with the taxonomy of Ctenuchini. We thank William Conner and Raleen Wil- son for providing specimens for DNA and morphological studies. We thank the following curators for access to their museum material: R. Holzenthal (Insect Museum Minnesota, IMSP), J. Y. Miller (Allyn Museum, FSMC), J. Minet (MNHP), M. Pogue (NMNH), and M. Scoble (BMNH). T. Perella assisted with dissections, and C. C Sasmey assisted with the molecular study. D. Vital provided illustrations. We thank M. DaCosta, W. Miller, D. Nickle, D. Smith, J. Zaspel, and anonymous reviewers for comments that improved this manuscript. The molecular study was supported in part by a University of Min- nesota, Undergraduate Research Opportunity Grant (UROP, UMN) to A. Carlson. Additional support was provided by NSF/DEB- 9306755 and NSF/DEB-9981416 (S. Weller) and a Dissertation Im- provement Grant NSF/DEB-9706192 (R. Simmons, S.Weller).

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Salkt, R. K., D. H. GELFAND, S. Stoffel, S.J. SCHARF, R. HIGUCHI, G. T. Horn, K. B. Muuuis & H. A. Erticu. 1988. Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase. Science 239:487-491.

SCHEFFER, S. J. 2000. Molecular evidence of cryptic species within Liriomyza huidobrensis (Diptera: Agromyzidae). J. Econ. En- tomol. 93:1146-1151.

SCHEFFER, S. J. & M. L. Lewis. 2001. Two nuclear genes confirm mitochondrial evidence of cryptic species within Liriomyza huidobrensis (Diptera: Agromyzidae). Ann. Entomol. Soc. Amer. 94:648-653.

SCHEFFER, S. J. & B. M. WEIGMANN. 2000. Molecular phylogenet- ics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): species limits, speciation, and dietary specialization. Mol. Phylo. Evol. 17:244-255.

Simmons, R. B. & S. J. WELLER. 2001. Utility and evolution of cytochrome b in insects. Molec. Phylog. and Evolution 20:196— 210.doi:10.1006/mpev.2001.0958

SIMON, C., F. Frati, A. Beckenbach, B. Crespt, H. Liu & P. FLOOK. 1994. Evolution, weighting and phylogenetic utility of mito- chondrial gene sequences, and a compilation of conserved PCR primers. Ann. Entomol. Soc. Am. 87:651—701.

SPERLING, F. A. H. & D. A. Hickey. 1994. Mitochondrial DNA se- quence variation in the Spruce Budworm species complex (Choristoneura: Lepidoptera). Mol. Biol. Evol. 11:656-665.

SworForb, D. L. 2000. “PAUP*—phylogenetic analysis using par- simony. (Version 4.0),” Sinauer, Sunderland.

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WELLER, S. J., R. B. Simmons, R. BoaDA & W. E. CONNER. 2000. Abdominal modifications occurring in wasp mimics of the Ctenuchine-Euchromiine clade (Lepidoptera: Arctiidae). Ann. Entomol. Soc. Am. 93:920-928.

Witson, R. 1999. Investigation of sound reception and production in Empyreuma pugione L. (Lepidoptera: Arctiidae). Unpub- lished MS Thesis (December 1999) Wake Forest University.

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Received 17 July 2002; revised and accepted 22 August 2003

Journal of the Lepidopterists’ Society 58(1), 2004, 33-43

THE BUTTERFLIES (PAPILIONOIDEA, HESPERIOIDEA) OF THE GALAPAGOS ISLANDS, ECUADOR: DISTRIBUTION, HOSTPLANTS AND BIOLOGY.

LAZARO ROQUE-ALBELO

Charles Darwin Research Station, Galapagos Islands, P.O. 17 01 3891, Quito, Ecuador, email: lazaro@fedarwin.org.ec. and Biodiversity and Ecological Processes Research Group, Cardiff School of Biosciences Cardiff Univ ersity, P.O. Box 915, Cardiff CF10 3TL, Wales, United Kingdom

ABSTRACT. The butterfly fauna of the Galapagos islands is reviewed. Ten species are recorded from the Archipelago. Leptotes parrha- sioides (Wallengren), Urbanus dorantes galapagensis (Williams), and Agraulis vanillae galapagensis (Holland) are endemic and the others are Neotropical faunal elements. A number of new island and host plant records are given.

Additional key words: _ flower visitation, phenology, Neotropical region.

The butterfly fauna of the Galapagos Islands was first discussed by Williams (1911). Six species were then recorded from the archipelago. Previous reports of Galapagos butterflies were published by Wallengren (1860, 1861), Butler (1877) and Holland (1889). Sub- sequent to Williams’ revision, little mention was made of the butterflies in the Galapagos until Beebe (1923) and Parkin et al. (1972) added two species of milk- weed butterflies, Danaus plexippus (Linnnaeus) and Danaus gilippus (Cramer), to the fauna. A literature- based checklist of the insects of Galapagos by Linsley and Usinger (1966) and Linsley (1977) listed eight species and indicated on which island each species oc- curred. During the last 33 years only two faunistic notes have been published. Onore and Mielke (1988) reported Calpodes ethlius (Stoll) and Roque et al. (1997) added Hemiargus ramon (Dognin). Additional information such as flower visits and host plant records were given by Linsley (1966), McMullen (1986, 1990, 1993), McMullen and Viderman (1994) and Roque (1998a, b). Finally, Peck et al. (1998) listed the intro- duced insect species that occur on the archipelago and included C. ethlius and H. ramon.

This paper reports new information on Galapagos butterflies gathered during 8 years of fieldwork on the Islands. The butterfly faunal of the archipelago in- cludes 10 species and subspecies, three of which are presumed to be endemic.

MATERIALS AND METHODS

From November 1994 to June 2002 the author con- ducted daytime observations and collections of butter- flies in the Galapagos Islands. A total of 387 specimens were sampled and identified, and observations on habitat associations and behavior of each species was noted. The methods included: a) field collections with a light mesh net, b) field observations of adult behay- ior and visits to food sources, c) collecting and rearing of immature stages.

Immatures were collected and reared in the labora- tory. Eggs and larvae were reared in plastic bags at room

temperature under the natural day-night photoperiod regime. Fresh leaves were provides at least ev ery other day. Pupae were kept moist until adults hatched.

The majority of this material was deposited in the Invertebrates Collection of the Charles Darwin Re- search Station (CDRS), Santa Cruz Island, Galapagos Islands. Some of the other specimens are in the fol- lowing collections: Pontificia Universidad Catélica del Ecuador (PUCE), Quito, Ecuador, Museo Ecuatori- ano de Ciencias Naturales (MECN), Quito, Ecuador and California Academy of Sciences (CAS), San Fran- cisco, USA. Additional records of distributions were obtained from the literature and unpublished data from Dr Bernard Landry, whose material is deposited in the Canadian National Collection (CNC), Ottawa, Canada.

The classification adopted here follows Ackery and Vane-Wright (1984) and Ackery et al. (1999). A brief description i is given for the immature stages of the en- demic species. The nomenclature of the plants follows McMullen (1999).

The following abbreviations were used: FW = fore wings, DFW = dorsal fore wings, VFW = ventral fore wings. HW = hind wings, DHW = dorsal hind wings, VHW = ventral hind wings. DW = dorsal side of the wings, VW = ventral side of the wings. The size of the butterflies in mm refers to their fore wing length.

RESULTS

Doubtful Records

Previous authors reported several species whose occurrence in the Galépagos has not been demon- strated convincingly. It is better to exclude these as probable misidentifications or mislabelling as is ex- plained below.

Actinote sp. (Nymphalidae): Daniel Fitter, a natu- ralist guide with the Galapagos National Park Service, sent a color photograph to the author for examination. The photograph, taken on Media Luna, Santa Cruz Is- land, was published in Fitter et al. (2001:115) and was

34

identified as Actinote sp. (Nymphalidae). It has not been collected or seen in the archipelago since the pic-

ture was taken. The establishment in the Galapagos of

this species will be only confirmed or refuted when ad- ditional material is found.

Hypolimmnas misippus (Linné), (Nymphalinae): The only notice of the occurrence of this species in the archipelago is from one specimen deposited in the American Museum of Natural History, New York. The specimen has the following label data: “Conway Bay, Indefatigable Is, Galapagos, March 15 1935, Crocker Exped’. It is likely that this is a mislabelling or if the record is valid it was a migrant individual. The species has never been seen or collected since.

Agraulis vanillae lucina (C. & R. Felder): This species was reported by Moreno et al. (1997). It is likely a misidentification of the Galapagos’s endemic sub- species Agraulis vanillae galapagensis (Holland, 1889).

Heteropia sp. (Hesperiidae): One male from Albe- marle [Isabela] was reported by Van Duzee (1933), al- though the author suggested that the specimen had been mislabeled and collected in Mexico.

Erynnis funeralis (Scudder & Burgess), (Hesperi- idae): One specimen (as Thanos persius (Scudder)) deposited in the CAS is said to have been collected in Chatham [San Cristdbal], 15 IV 1932 by M Willows Jr. However, Van Duzee (1933) suggested the specimen was mislabelled.

Leptotes trigemmatus (Butler), (Lycaenidae): Larsen (1991) mentioned this species from the Gala- pagos Islands but it is likely a misidentification of the sister taxon Leptotes parrhasioides (Wallengren), as noted by Balint and Johnson (1995).

Phoebis agarithe (Boisduval) (Pieridae): Moreno et al. (1997) reported this species from the archipelago. These authors probably misidentified Phoebis sennae marcellina (Cramer) as this species which is superfi- cially similar in size and color. P. agarithe has never been collected on the Galapagos Islands.

SPECIES ACCOUNTS HESPERIIDAE Pyrginae Urbanus dorantes zalapagensis (Williams 1911) “Galapagos Skipper” (Fig. 1) Eudamus galapagensis Williams 1911:303; Draudt 1921:853; Beebe 1923:55; Van Duzee 1933:144. Urbanus dorantes galapagensis; Evans 1952:93; Linsley & Usinger 1966:157; Parkin et al. 1972:103. Urbanus galapagensis Williams & Hoyward 1944:93. Diagnosis. FW = 15-20 mm. This is the only tailed skipper of the archipelago. FW dark brown with

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translucent spots and a pattern of lilac and brown bands in VHW. In some specimens, the hyaline spots are smaller. The lilac bands on the VW tend to be pale with lighter brown or yellowish scales. The sexes are stivatllere but males have coastal folds.

Geographical distribution. Reported from Bal- tra, Edén, Espafiola, Floreana, Genovesa, Isabela, Pinzon, San Cristébal, Santa Cruz and Santiago (Lins- ley & Usinger 1966). Additional records include: Fer- nandina, Pinta and Rabida.

Natural history. Adults were common in open ar- eas, roadsides, and forests from sea level to the highest elevations. The flight is erratic and low to the ground. Both sexes were seen visiting flowers. Oviposition was observed on leaves and flower buds of various Legu- minosae. Eggs greenish laid singly on Phaseolus atro- purpureus D.C., P. mollis Hook, Desmodium incanum DC and Galactia striata (Jacq). Caterpillar green with the head and prothoracic shield black. In the last instar the body is brown. Pupa pale brown speckled with darker brown.

Flowers visited. Acanthaceae: Justicia galapagana Lindau; Asteraceae: Darwiniothamnus tenuifolius (Hook. f.) Harling, Scalesia cordata Stewart, Scalesia microcephala Robins, Scalesia villosa Stewart, Scalesia retroflexa Hemsley; Boraginaceae: Cordia leucophlyc- tis Hook, C. lutea Lam.; Convolvulaceae: Merremia aegyptica (L.) Urban, Leguminosae: Acacia insulae- iacobi Riley, P. mollis; Malvaceae: Sida rhombifolia L.; Plumbaginaceae: Plumbago scandens L.; Rubiaceae: Diodia radula Cham.& Schlecht; Rutaceae: Citrus limon (L.) Burm; Solanaceae: Lycopersicon chesmanii Riley, Physalis pubescens L.; Sterculiaceae: Waltheria ovata Cav.; Verbenaceae: Lantana camara L., Lantana peduncularis Anderss, and Lippia strigulosa Mart & Gal.

Flight period. Multivoltine with a peak in January and February.

Remarks. The holotype is in the California Academy of Sciences, San Francisco, USA (CAS): it bears the fol- lowing label information: Galapagos Is, Chatham I. [San Cristobal], X 15 06, Coll. By F. X. Williams, Califomia Academy of Sciences Type No. 3209. A specimen de- posited on CAS has the following label information: Puerto Vallarta, Jalisco, Mexico, VIII 21 32, M Willow Jr coll, Exp 1932. This could be a mislabelling. Evans (1952) treated it as a subspecies of Urbanus dorantes (Stoll, 1790) compared to Urbanus dorantes santiago (Lucas, 1856) from Cuba and U. dorantes dorantes from mainland Ecuador, U. dorantes galapagensis is smaller and has shorter hindwing tails. The male valve is much less dentate and the arms of harpe have few spines. The genus Urbanus Hiibner 1807, is used provisionally here until revisionary work is undertaken. According to S.

VOLUME 58, NUMBER 1 35

7 | ; 8 |

Fics. 1-8. Galapagos butterflies species. 1, Urbanus galapagensis (DW and VW); 2, Calpodes ethlius (DW); 3, Male of Leptotes parrha- sioides (DW and VW); 4, Male of Hemiargus ramon (DW); 5, Male of Hemiargus ramon (VW); 6, Female of Hemiargus ramon (DW); 7, Fe- male of Phoebis sennae marcellina (DW); 8, Male of Phoebis sennae marcellina (DW).

36

Steinhauser (pers. com.) the species may belong in a separate, undescribe genus.

Material studied. (706 and 16°). ECUADOR, Galapagos. ES- PANOLA: Hood, 15 V 1899, [R. E. Snodgrass, E. Heller] (CAS) (1d). Bahia Manzanillo, 24 IV 1992, B. L[andry] (CNC), (2¢). FER- NANDINA: 1 IX 1966, (CDRS) (12), Cueva Norte, Arid zone, 20 VI 1998, L. Roque] (CDRS) (16). FLOREANA: Arid zone, 130 m, 17 IV 1996, L. R (CDRS) (8¢ and 32). Cerro Pajas, Scalesia forest, 320 m, 18 IV 1996, L. R (CDRS) (6c). Cerro Pajas 520 m, 19 IV 1996, L. R (CDRS) (9¢). Charles, 5 X 1899, [R. E. Snodgrass, E. Heller] (CAS) (5d i ISABELA: Tagus Cove, II] 1906, Alotype, FX Williams] (CAS) (12). Cowley Mt, VIII 1906, F. X. W (CAS) (3 3) Albemarle, 27 IV oe M. Willows (CAS) (1¢); 28 IV 1932, M. Wil lows (CAS) (12). Albemarle, 12 VI 1899, [R. E. Snodgrass, E. Heller] (CAS) (12). Albemarle Iguana Cove, 21 V 1932, M. Willows cS AS) (22). V[olean] A[lcedo| | 800 m., 2 IV 1996, e R (CDRS) (5d

). V com N[egra], 2 III 1989, S. B. Peck (CNC) (1¢). VSN, 6 IV Ate L. R (CDRS) (1d 12). VSN, Las merce ne 120m, 24 IV 1996, L. R (CDRS) (1¢ 72). Sierra Negra pampas ferns zone, 24 IV 1996, L. R (CDRS) (12). Santo Tomas, 20 TIT 1995, P. Delgado (CDRS) (32); 6 IV 1996, L. R (CDRS) (2¢); 23 IV 1996, L. R (CDRS) (1¢); 23 TV 1996, Cae (1d). V W[olf], 1200 m, Scalesia forest, 19 IV 1996 (CDRS) (7¢ 42). Playa Tortuga Negra, 22 VI 1998, L R (CDRS) (12). PINTA: Abingdon, 24 VI 1899, [R. E. Snodgrass, E. Heller], (CAS) (12). PINZON: Pl: aya escondida, 20 TV 2002, L. R & B. Landry (CDRS) (26 39). RABIDA: Sendero turistico, 13 III 1998, L. R, (CDRS) (1¢). SANTA CRUZ: Horne- man farm 220m, 18 III 1964, (1d); 3 V 1964, (22), D. Q. Cavagnaro, (CAS). Puerto Ayora, 17 I 1989, B. L (CNC) i ‘); 8 XI 1995, A. Herrera (CDRS) (1¢); 29 X 1995, L. R, V. Cruz (CDRS) (1¢). 4 km N Puerto Ayora, 20 I 1989, B. L (MECN) (1¢). Tortuga Reserve W Santa Rosa, 6 V 1989, S. B. Peck (CNC) (12); 6 II 1989, B. Landry (CDRS) (12). Cerro Crocker, 9 IT 1996, L. R (CDRS) (1d), Bella- vista, 13 II 1999, reared from leaves D. canum, L. R (CDRS) (4¢ 19); Punta Nufiez, 29 VII 2002, R. eee (CDRS) (12). SAN CRISTOBAL: Chatam, X aoe F. X. W, (CAS) (3d): 1 X 1906, Holotype, F. X. W (CAS) (1d); 23 V Hee. [R. E. Snodgrass, E. Heller], (CAS) (6¢ 19); 15 ey 1932, (3d); 17 IV 1932, (1d); 18 IV 1932, (23), M. Willows, (CAS). Puerto Baquerizo, 17 II 1989, S. B. Peck (CNC) (12). Airport 30 m, 6 TIT 1996, L. R (CDRS) (6 $99). SANTIAGO: James, 21 IV 1899, [R. E. Snodgrass, E. Heller], (CAS) (22). Central 700 m, 9 IV 1992, B. L (CNC) (12).

Hesperiinae Calpodes ethlius (Stoll, 1782) “C (Fig. 2)

Papilio ethlius Stoll 1782:212. Calpodes ethlius, Onore & Mielke 1988:619; Peck et al. 1998:227.

anna Skipper”

Diagnosis. FW = 19.5-24 mm. The Canna Skipper isa large brown species with translucent spots on the wings. Diagnostic features include a linear row of four spots on the HW and the absence of a tail. Specimens collected in the Galapagos are similar in shape and size to mainland specimens.

Geographical distribution. This species is widely distributed from Florida to Argentina including the West Indies. In the Galapagos Islands it has been re- ported previously from Isabela (Onore & Mielke 1988) and here from Santa Cruz Island.

Natural history. Calpodes ethlius is a powerful flyer that is most active at dawn and dusk, but is also seen at other times of the day. The species is most

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abundant in urban areas and gardens but it has the ability to find isolated hosts inside the forests. Eggs are pale green and laid singly on the leaves of Canna edulis Kerr and Canna lutea Mill (Cannaceae), which have been introduced to the Archipelago.

Flowers visited. Verbenaceae: Clerodendrum molle HBK.

Flight period. Specimens have been collected in February, April, May and July.

Remarks. Peck et al. (1998) listed this large skip- per as recently introduced in the archipelago, and it is a known migrant (Ferguson et al. 1991).

Material studied. (5d and 1°). ECUADOR, Galapagos. IS- ABELA: VSN, 1 km W Santo Tomas, 12 V 1996, ex larvae, L. R (CDRS) (1d). SANTA CRUZ: Road to Baltra, 11 VI 1981, Y. Lu- bin, (CDRS) (1¢). Puerto Ayora, 24 VII 1996, (CDRS) (32); 18 II 1998, (CDRS) (12); 22 IV 1998, L. R (CDRS) (1¢)

PIERIDAE

Coliadinae Phoebis sennae marcellina (Cramer, 1779) “Sulfur Butterfly” (Figs. 7, 8) Papilio marcellina Cramer 1779:103. Callidryas eubele; Holland 1889:195; Williams 1911:296; Beebe 1923:55. Catopsila eubele; Van Duzee 1933:140. Phoebis sennae; Linsley & Usinger 1966:156; Parkin et al.

1972:103.

Phoebis sennae marcellina; Moreno et al. 1997:146.

Diagnosis. FW = 24-36 mm. Easily recognized, it is the only yellow butterfly on the Galapagos Islands. Male: bright yellow. Female: yellow with black spots along the outer margins of the wings and a black spot with a pale center at the end of the forewing cell above. Galapagos populations are very variable in color pattern and size, but are identical with mainland spec- imens in genitalic features.

Geographical distribution. Phoebis sennae mar- cellina flies from México to Uruguay (D’Abrera 1981). In the Galapagos Islands the species has been previ- ously reported from Baltra, Daphne, Edén, Fernand- ina, Floreana, Isabela, Pinta, Santiago, San Crist6bal, and Santa Cruz (Linsley & Usinger 1966) and is here reported from Marchena.

Natural history. One of the most common species on the Galapagos Islands. Adults occur in open habi- tats such as beaches, dry forests and roadsides from sea level to 900 m. Both sexes have been observed vis- iting flowers and puddles. The flight is fast and erratic. Beebe (1924) reported a southeast migration of this species from Santiago and Isabela to Santa Cruz, and individuals have been observed commonly flying be- tween islands. Williams (1911) reported Senna picta G. Don as a host plant in Galapagos. Females have

VOLUME 58, NUMBER 1

been observed laying eggs in Senna tora L., Senna bi- capsularis L., and Senna occidentalis L. (Fabaceae).

Flowers visited. Acanthaceae: J. galapagana; Apocynaceae: Catharanthus roseus (L.) G. Don; Aster- aceae: Bidens pilosa L.; Boraginaceae: C. leucophlyctis, C. lutea, Heliotropium angiospermum Murr., Tourne- fortia rufo-sericea Hook. f.; Cactaceae: Opuntia insu- laris Stewart; Convolvulaceae: Ipomoea pes-caprae (L.) R. Br., Ipomoea nil (L.) Roth; Leguminosae: S. picta; Malvaceae: Gossypium barbadense var. darwinii (Watt.) Hutch, Hibiscus rosa-sinensis L., S. rhombifo- lia; Nyctaginaceae: Bougainvillea spectabilis Willd, Plumbaginaceae: P. scandens; Rubiaceae: D. radula; Verbenaceae: L. camara, Verbena litoralis HBK.

Flight period. Multivoltine. Specimens have been seen throughout the year.

Material studied. (326 and 102). ECUADOR, Galapagos. FERNANDINA: Cabo Douglas, 15 IV 1999, P. Polo (CDRS) (1). ISABELA: VA, Pega-pega, 6 IV 1999, L. R{oque] (CDRS) (1¢). Puerto Villamil, 2 III 1989, B. L{andry] (CNC) (1¢). Arid Zone, 7 III-1989, S. B. Peck (CNC) (1d). VSN, 6 IV 1996, L. R (CDRS) (5d and 22). VSN, Santo Tomas 300 m, 26 IV 1996, L. R (CDRS) (8¢). VSN, Pampas, 24 IV 1996, L. R (CDRS) (1¢ and 32). VSN, La Bo- canilla 1000 m, 28 IV 1996, L. R (CDRS) (2d). VSN, Alemania 350 m, 29 IV 1996, L. R (CDRS) (2¢). Playa Tortuga Negra, 22 VI 1998, L. R (CDRS) (3¢). MARCHENA: Playa } Negra, 30. TV 2000, L. R, (CDRS) (1d y: SAN CRISTOBAL: Puerto Baquerizo, 17 II 1989, B. L (CDRS) (2¢): El Porvenir, 15 III 1996, L. R (CDRS) (1¢ and 52). SANTA CRUZ: Santa Cruz, 8 VIII 1990, C.K McMullen (CDRS) (13). SANTIAGO: Los Jaboncillos, XII 1974, (CDRS) (1d); Es- pumilla, 17 III 1998, L. R (CDRS) (1¢).

LYCAENIDAE

Polyommatinae Leptotes parrhasioides (Wallengren 1860) “Galapagos Blue Butterfly” (Fig. 3) Lycaena parrhasioides Wallengren 1860:37; 1861:355. Cupido parrhasioides; Williams 1911:300, Beebe 1923:55; Van Duzee 1933:143; Moreno et al. 1997:158.

Leptotes parrhasioides; Bridges 1988:I 268, Linsley & Usinger 1966:157; Balint & Jonhson 1995:9.

Diagnosis. This species and the following are the only lycaenid butterflies in the Galapagos Islands. Both are similar, but the wing pattern and genitalia are distinctive. FW = 8-12.5 mm. Male: DFW and DHW violet blue with two or three black anal spots. VW ground color grayish with a pattern of white markings. VHW with three or four marginal spots, black in the middle, and filled with metallic blue. Female: similar to male but DW ground brown or blue.

Geographical distribution. Endemic to the Gala- pagos Islands, and reported from the islands of Baltra, Edén, Espafiola, Fernandina, Floreana, Isabela, Pinzon, San Cristobal, Santa Cruz, Santiago (Linsley & Usinger 1966). It is reported here from Marchena, Pinta and Rabida.

~l

eo

Natural history. Common in the arid zone of the islands. The flight is slow and close to the ground, but may also be rapid and erratic. Both sexes have been seen visiting flowers and were often observed on muddy puddles early in the morning or after rainfall. The pale green eggs are laid singly on the flowers, young fruits, or young leaves of host plants. Williams (1911) reported Cardiospermum halicacabum L. as a host plant and indicated Cardiospermum gala- pageium B. L. Rob. & Greenm (Sapindaceae) as another probable food plant. New hostplants records include: P. scandes, Crotalaria incana L. and Rhyn- chosia minima (L.) DC. (Leguminosae). The larvae are variable, either green or red with a pattern of dark dorsal marks, depending on the host. The body is cov- ered by short white hairs. Pupae are greenish with brown markings.

Flowers visited. Acanthaceae: J. galapagana; Asteraceae: Darwiniothamnus lancifolius (Hook. f.) Harling; Boraginaceae: H. curassavicum, Tournefortia valoda ya HBK, T. rufo-sericeae; Cucurbitaceae: Momordica charantia L.; Cyperaceae: Cyperus ligu- laris L.; Leguminosae: Vigna luteola (Jacq.) Benth.; Linaceae: Linum cratericola Eliass; Lythraceae: Cuphea racemosa (L. f.) Spreng.; Malvaceae: S. rhombifolia, Plumbaginaceae: P. scandens; Nyctagi- naceae: Commicarpus tuberosus (Lam.) Standl.; Solanaceae: L. cheesmanii; Zygophyllaceae: Tribulus cistoides L.

Flight period. The species is multivoltine.

Remarks. Wallengren (1860) described this en- demic species from at least one male and one female [locality not specified] collected in May 1852 on the Galapagos. However, in 1861 Wallengren redescribed the species with additional material labelled “Puna” [gulf of Guayaquil] and “San Jose island” [Panama]. Balint and Jonhson (1995) studied a specimen labeled “type” in the Naturhistoriska Riksmuseet, Stockholm, Sweden (NRS). They stated that this specimen be- longs to the sister species Leptotes trigemmatus (But- ler) and that the specimens involved in the redescrip- tion (Wallengren 1861) were incorrectly associated with the original series from the Galapagos. The real type of L. parrhasioides was not examined, although it is probably located in NRS.

Material studied. (526 and 122). ECUADOR Galapagos. ES- PANOLA: Bahia Manzanillo, 25 IV 1992, B. L (CNC) (1¢). FER- NANDINA: Narborough, 16 IV 1906, F X W[{illiams], (CDRS) (1°). Near to summit 4800ft, 5 IT 1964, D. Q. Cavagnaro, (CAS) (3°). Cabo Douglas, 21 VI 1998, L. R[oque] and C. Clauston], (CDRS) (3¢ and 2°). Campamento Perez South Punta Espinoza, 18 VI 1998, L. R and C. C, (CDRS) (7¢ and 42). Cueva norte, 20 VI 1998, L. R and C. C, (CDRS) (4¢ and 2°). North Side 300m, 12 I 2002, L. R and C. C, (CDRS) (12). Zona de vegetacion, 20 VI 1996, L R, (CDRS) (4¢ and 32). FLOREANA: Charles, 10 X 1905, F X W, (CAS) (22): 23-31 V

1906, F X W, (CAS) (19); 14 VI 1906, FXW, (CAS) (2¢). Zona costera, 2 TV 1995, L. R and V. Cruz, (CDRS) (1¢). Scalesia forest, 4 III 1995, L. R and V. Cruz (CDRS) (12¢ and 42). Arid zone 130m, 4 III 1995, in H. curassavicum flowers (3¢); in C. molle flowers (1¢), L. R and V. Cruz (CDRS). Road to the highland, 4 III 1995, L. R and V. Cruz (CDRS) (3¢). Parte alta, 4 III 1995, L. R and V. Cruz (CDRS) (1d). Cerro Alieri 340 m, 21 XII 1999, in Linum cratericola flowers, L. R, (CDRS) (1d). ISABELA: Albemarle Banks Bay, 16 X 1906, F X W, (CAS) (2d). Albemarle Tagus Cove, 22 III 1906, F X W, (CAS) (1d). V[olean] S[ierra] N[egra] 1000 m, 1 III 1989, B. L (CNC) (12). V[olean] W[olf], Scalesia forest 1700 m, 19 V 1996, L. R (CDRS) (4d). V W, Upper arid zone, 1250 m, 20 IV 1996, L. R (CDRS) (132). Playa Tortuga Negra, 22 VI 1998, L. R, (CDRS) (8

and 7°). PINTA: Playa Ibetson, 14 HI 1992, B. L (CNC) (1¢ and 19); reared from leaves R. minima, 14 III 1992, B. L (CNC) (1¢). 400 m, 17 III 1992, B. L (CNC) (22). PINTA: Abingdon green zone, 10 IV 1906, F X W, (CAS) (12). PINZON: Summit & upper caldera areas, 7 IL 1964, D. Q. Cavagnaro, (CAS) (1¢) a ey AGO: James bay low altitud, VIII 1932, M. Willows, (CAS) _ SANTA CRUZ: Inde-

fatigable, 8 VI 1932, M. Willows, ae aaa 12). Academy Bay CDRS, 5 II 1964, R. O. Schuster, (CAS) (1¢); 25 II 1964, D. Q. Cav- agnaro and R. O. Schuster, (CAS) (1¢). aud Zone, asociated to Castela galapageia, 19 I 1989, B. L (CNC) (2°). Transition zone, 20 I 1989, B. L (CNC) (1¢). SAN CRISTOBAL: Chatam Sapho cove, 9 II 1906, F X W, (CAS) (1¢)

Hemiargus ramon (Dognin, 1887) “Ramon’s Blue Butterfly” (Figs. 4-6) Lycaena ramon Dognin 1887:23. Hemiargus ramon; Roque et al. 1997:29; Peck et al. 1998:227.

Diagnosis. FW = 8.2-11.6 mm. Resembles L. par- rhasioides, with which it flies, but H. ramon has two black dots in the VHW inner costal margin while L. parrhasioides has none. Male: DW and DHW violet blue, the outer margins thinly bordered with white. Female: DW and DHW dark with only a touch of blue at the wing base.

Geographical distribution. Distributed on the western slopes of the Andes from northern Chile to the south of Ecuador. This species was listed by Peck et al. (1998) as introduced in the Archipelago. It has been reported from the Galapagos Islands of Baltra, Espafiola, Floreana, Isabela, San Crist6bal, Santa Cruz and Santiago (Roque et al. 1998) and is here reported from Fernandina, Pinz6én, Santa Fé, Seymour Norte and Rabida.

Natural history. Common from the low arid zones to the highest elevations. Roque et al. (1997) reported seven leguminous plants as host of the species (Acacia macracantha Willd, S. tora, Neptunia plena Benth, P. atropurpureus, Prosopis juliflora D.C., Tephrosia de- cumbens (Benth), and one species of Cucurbitaceae: Cucumis dipsaceus Her. and one species of Oxali- daceae is reported here: Oxalis dombeyi A. St-Hil.

Flowers visited. Asteraceae: Scalesia gordilloi Hamann & Wium Anders., S. pedunculata; Boragi- naceae: T. rufo-sericeae; Leguminosae: T. decumbens.

Flight period. Observed during all months of the

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

year, but is most common during the rainy season (Jan- uary to April).

Material studied. (42¢ and 12°). ECUADOR Galapagos. ESPANOLA: Bahia Manzanillo, 24 IV 1992, (1¢); 25 IV 1999 (1¢), B. Landry, (CNC). FERNANDINA: Campamento Perez south Punta Espinoza, 18 VI 1998, L. Rf{oque] and C. Clauston] (CDRS) (1d and 29); Cabo Douglas, 21 VI 1998, L. R and C. C, (CDRS) (1¢). Zona de vegetacién, 20 VI 1998, L. R and C. C, (CDRS) (12), FLOREANA: Zona Costera, 2 [TV 1995, L. R and V. C[ruz] (CDRS) (8¢ and 52). High arid zone 150m, 4 III 1995, L. R and V. C (CDRS) (5d). Road to Highland, 4 IIT 1995, L. R and

5. (CDRS) (10¢ and 3°). Parte media, 4 III 1995, in He- liotropium curassavicum flowers, L. R and V. C, (CDRS) (5¢). IS- ABELA: V[olcan] S[ierra] N[egra], 24 TV 1996, L. R (CDRS) (24 and 2°). VS N, Las Merceditas 120m, 24 IV 1996, L. R (CDRS) (12). VS N,2km W Puerto Villamil, 3 VI 2000, C. Covell (CDRS) (6d and 29). aes Playa escondida, 20 VI 2002, L. R and B. L, (CDRS) (3¢ and 3°). SANTA CRUZ: Arid Zone, 8 IX 1999, L. R and C. Covell (CDRS) (22 and 4¢); 19 I 1989, B. L, (CNC) (3¢ and 39).

NYMPHALIDAE Danainae Danaus gilippus thersippus (Bates 1863) “Queen Butterfly”

(Fig. 11) Papilio thersippus Bates 1863. Danaus gilippus; Parkin et al. 1972:103; Linsley 1977:29. Danaus gilippus thersippus; Ackery & Vane-Wright 1984:208.

Diagnosis. FW = 36-41 mm. Easily distinguished from D. plexippus L. by not having the black inner border, and the black veins on the DFW. Males have a black androconial patch on the DHW. Females are similar to males but without an androconial patch.

Geographical distribution. Danaus gilippus ther- sippus is distributed from the Southern United States to Panama and is native to the Galapagos Islands. The species has been reported from San Cristobal (Parkin et al. 1972), and is here reported from Baltra, Flore- ana, Isabela, Santa Cruz, Santiago, Seymour Norte and Rabida.

Natural history. The queen butterfly is primarily an inhabitant of the arid zones from sea level to 400 m, but its distribution is limited by the occurrence of its foodplants. Adults have been observed flying slowly and frequently visiting flowers in open areas. The pale green eggs are laid singly on young and old leaves of the endemic vine Sarcostenma angustissima R. W. Holm. The other milkweed that occurs on the islands, Asclepias curassavica L., apparently is used as an al- ternative food plant.

Flowers visited. Asclepiadaceae: A. curassavica, S. angustissima; Asteraceae: D. tenuifolius; Boragi- naceae: H. angiospermum; Solanaceae: P. pubescens; Verbenaceae: L. camara.

Flight period. All available records are from the rainy season.

VOLUME 58, NUMBER 1 39

i153, 16

Fics. 9-16. 9, Agraulis vanillae galapagensis (DW); 10, Agraulis vanillae galapagensis (VW); 11, Danaus gilippus thersippus (DW); 12, Danaus plexippus megalippe (DW); 13, Vanessa virginiensis (DW); 14, Vanessa virginiensis (VW); 15, Vanessa carye (DW); 16, Vanessa carye (VW).

40

Remarks. Ackery and Vane-Wright (1984) sug- gested that this species could have been introduced ona California to the islands in recent times. Based on its current distribution and the well-established dis- persal potential, and taking into consideration the dates of discovery of the butterfly and its host plants, it is reasonable to assume that D. g. thersippus arrived in the Archipelago through Hee dispersal mechanisms and was not introduced by humans.

Material studied. (23¢ and 9°) ECUADOR, Galapagos. FER- NANDINA:, Cueva Norte, L. R[oque], (CDRS) (26 and 12). Cam- pamento Perez south Punta Espinoza, 18 VI 1998, L. R and C. Causton, (CDRS) (1d). FLOREANA: Cerro Pajas, 20 IV 1996, L. R (CDRS) (12). Arid zone, near to Black beach, 22 IV 1996, L. R (CDRS) (2d). C. Cruz farm, 24 VI 1996, L. R (CDRS) (4¢). Asilo de la Paz 342m, 23 IV 1996, L. R (CDRS) (49 and 42). ISABELA: Playa Tortuga Negra, 22 IV 1998, L. R (C DRS) (8 rae 32). Volcan Sierra Negra Nlemrania 350 m, 29 IV 1996, L. R, (CDRS) (10¢ and 62). Vol- can Sierra Negra pampas, 24 IV 1996, L. R, (CDRS) (12). SAN

CRISTOBAL: Puerto Baquerizo, 17 HT 1989, B. L, (CDRS) (1¢); (MECN) (1¢).

Danaus plexippus megalippe (Hiibner, [1826]) “The Monarch” (Fig. 12) Anosia megalippe Hiibner [1826}:2, pl. 7, fig. Danais plexippus; Beebe 1923:55. Danaus plexippus; Linsley & Usinger 1966:157; Roque 1998a:9, b:10. Danaus plexippus megalippe; Ackery & Vane-Wright 1984:201.

Diagnosis. FW = 35-50 mm. This species is easily distinguished from D. gilippus by having the DFW apex boldly black, a black inner border in DFW and prominent black veins. Males have a black androconial patch on the DFW. Females are similar to males but much darker.

Geographical distribution. The Monarch is na- tive in the Galapagos Islands and reported from Flore- ana, San Cristébal and Santa Cruz. The status of this species has been discussed in some detail by Roque (1998a). Danaus plexippus megalippe is distributed from Nicaragua to Pertti including the North east of Brazil, Venezuela, French Guia Surinam, Guyana, Trinidad, Tobago and some islands of the West Indies (Ackery & Vane-Wright 1984).

Natural history. Common in open areas such as fields, roadsides and disturbed sites. In general, the lo- calities where the species occurs are mesic, but there are a few records in arid zones. Galapagos populations do not migrate. The adults fly slowly and visit flowers. Females have been observed ovipositing on the milk- weeds A. curassavica and S. angustissima, but the lat- ter is used as an alternative host plant.

Flowers visited. Asclepiadacea: A. curassavica; Malvaceae: H. rosa-sinensis: Verbenaceae: L. camara, V. litoralis.

Flight period. Adults commonly occur only in the

Jc JURNAL OF THE LEPIDOPTERISTS’ SOCIETY

rainy season, but are observed sometimes in the dry season.

Material studied. (13¢ and 5°). ECUADOR, Gal: dpagos. FLO- REANA: Cerro Pajas, 20 IV 1996, I ase (CDRS) (1¢ and 19). fa zone, near to Black beach, 22 Iv 1996, L. R, (CDRS) (4¢ and

). Finca de C. Cruz, es VI 1996, L. R (C DRS) ) (12). Asilo de la Paz a m, 23 IV 1996, L. R, (CDRS) (4¢). SAN CRISTOBAL: Puerto Baquerizo, 17 II 1989. B. alan (ORS (1d); (CNC) (19); (MECN) (12). SANTA CRUZ: El cascajo, IT 1999, L. R, (CDRS)

). Salasaca, 22 VIII 2000, R. Oquendo, (CDRS) (1¢).

Heliconiinae Agraulis vanillae galapagensis (Holland 1889) “Galapagos Fritillary” (Figs. 9, 10) Agraulis vanillae galapagensis Holland 1889:194; Williams 1911:298; Beebe 1923:55; Van Duzee 1933:141. Dione vanillae galapagensis; Stichel 1938:92; Michener 1942:4; Linsley & Usinger 1966:156.

Diagnosis. FW = 19-29 mm. This species is distin- guished by its silver markings on the VW. The females are paler than males.

Geographical distribution. Endemic to the Gala- pagos Islands. It has been collected on Baltra, Edén, Fernandina, Floreana, Isabela, Pinta, San Cristobal, Santa Cruz, and Santiago (Linsley & Usinger 1966).

Natural history. This is a common species on the larger Galapagos Islands. Adults have been observed in open areas and forests from sea level to the high- lands. Beebe (1924) cited an interesting migration along the West Coast of Isabela Island. Adults usually fly slow, low to the ground and frequently have been seen visiting flowers. Williams (1911) mentioned species of the genus Passiflora L. as food plants of this fritillary in the Archipelago, but he did not mention which. Oviposition was observed on Passiflora suberosa L. and Passiflora tridactylites Hook. (Passi- floraceae). The eggs were laid singly on young leaves. The larvae are black with green longitudinal stripes. The head and body are covered with long black spines.

Flowers visited. Acanthaceae: Blechum brownei Juss. f., J. galapagana; Asclepiadaceae: S. angustis- sima; Boraginaceae: Croton scouleri Hook. f.; Can- naceae: Commelina diffusa Burdm. f.; Malvaceae: S. hederifolia Cav., S. paniculata L., S. salviifolia Presl.; Rubiaceae: D. radula; Rutaceae: C. limon; Sterculi- aceae: Waltheria ovata Cav.; Verbenaceae: L. camara, Stachytarpheta cayennensis (Rich.) M.Vahl.

Flight period. All specimens were collected in the rainy season but individuals were observed during all months of the year except August and September.

Remarks. This species has been treated by sev- eral authors as a subspecies of the widely distributed American species Agraulis vanillae. Holland (1889) and Williams (1911) rerered that the Galapagos taxon

VOLUME 58, NUMBER 1

should be raised to specific status based on several differences in the wing pattern. Michener (1942) rec- ognized the differences in the wing maculation and size, and mentioned some variation in the male geni- talic valvae. Specimens studied here were collected on several localities of the Archipelago.The male gen- italia are generally similar to those of mainland speci- mens of A. vanillae but differ consistently in the valva. The apex of the valva is more slender and with fewer denticules in A. v. galapagensis. The female genitalia are similar in both forms. Agraulis vanillae galapa- gensis can be separated from the other described sub- species by the smaller size, by the darker and more fuscous color of the basal half of the wings dorsally, by the stronger black markings on the wings, and by the absence of white dots in the black spots of the DFW discal cell. The holotype, collected on Chatham [San Crist6bal], is deposited in the National Museum of Natural History, Washington, DC (USNM). The iden- tity of this species will remain questionable until a re- vision of the genus makes it more definite.

Material studied. (14¢ and 512). ECUADOR, Galapagos. FERNANDINA: Campamento Perez south Punta Espinoza, 18 VI 1998, L. R[oque] and C. Causton, (CDRS) (1d). FLOREANA: Arid zone 130m, 17 IV 1996, L. R, (CDRS) (1d). Cerro Pajas, Scalesia forest, 18 IV 1996, (94 and 59); 19 IV 1996, L. R, (12); 22 IV 1996, L. R (CDRS) (7¢ and 29). ISABELA: V[olean] S[ierra] N[egra], Santo Tomas, 6 IV 1996, (12): 26 IV 1996, L. R, (CDRS) (1d). VSN, bosque de los nifios, 20 HII 1995, L. R, (CDRS) (1d). VS N, pampas 1000m, 1 II 1989, B. L{andry] (CNC) (12). V SN, la bocanilla 1000m, 28 IV 1996, L. R, (CDRS) (2¢ and 12). VS N, Las mercedi- tas, 30 IV 1996, L. R, (CDRS) (2¢ and 42). Volcan Wolf, upper arid zone 1700 m, 19 V 1996, L. R, (CDRS) (13d and 42). PINTA: 400 m, 17 II 1992, B. L, (CNC) (42). SANTIAGO: Playa Espumilla, 4 IV 1992, B. L, (CNC) (1d). SANTA CRUZ: CDRS, 22 XI 1963, (CDRS) (1¢). Tortugas reserve west of Santa Rosa, 6 II 1989, B. L, (CDRS) (16); (MECN) (1d).

Nymphalinae Vanessa virginiensis (Drury 1773) “Painted Lady” (Figs. 13, 14) Nymphalis cardui virginiensis Drury 1773:1. Pyrameis huntera; Williams 1911:299. Vanessa virginiensis; Linsley & Usinger 1966:156.

Diagnosis. FW = 21.7-29 mm. Differs from V. carye by the presence of two blue-centered ocular markings on the VHW. Males and females are similar.

Geographical distribution. This cosmopolitan species was previously reported from Isabela (Volca- noes Darwin and Sierra Negra) (Williams 1911). On Isabela, the species was found at those localities as well as on voleanoes Alcedo, Cerro Azul and Wolf. Other new records include Santa Cruz and Santiago and it probably also inhabits other large islands. The species is distributed from southern Canada to the mountainous region of Colombia, including the West

4]

Indies, Hawaii, Azores, Madeira, the Canary Islands. and occasionally Europe (DeVries 1987).

Natural history. This species has only been ob- served in the open grassy areas of the highlands. All specimens were collected above 900 m in the pampa zone. Although it flies usually under bright and sunny conditions, we have seen it under overcast conditions and even during light rain (Volcén Alcedo, 23.X.1998). Adults fly fast and rest on the ground, flying short dis- tances when disturbed. Eggs, larvae and pupae were found on the leaves of Gnaphalium purpureum L. (Asteraceae) on the islands of Isabela and Santiago. The plant can be found on more islands (Lawesson et al. 1987). Vanessa virginiensis has resident populations in the Galapagos Islands.

Flowers visited. Rubiaceae: Borreria laevis (Lam.) Griseb.

Flight period. Specimens have been collected or observed from January to June.

Material studied. (154 and 42). ISABELA: Albemarle, 20-31 VIII 1906, F X Williams, (CAS) (1d). V[oleén] S[ierra] N[egra], rim northwest side, 4 III 1989, B. Llandry], (CDRS) (1¢); (MECN) (1¢). VSN, 1000 m, 4 III 1989, S. B. Peck (CNC) (12). V S N, La Bo- canilla 1000 m, 28 IV 1996, L. R{oque], (CDRS) (1¢ and 42). VS N, Las Merceditas, 6 IV 1996, L. R, (CDRS) (1¢). Volcan Wolf, high arid zone, 1250 m, 21 V 1996, L. R, (CDRS) (43). Volcan Alcedo, top 1100 m, 24 IV 1998, L. R (24 and 42). SANTA CRUZ: CDRS, 10 VI 1965, (CDRS) (1¢). SANTIAGO: Los Jaboncillos 900 m, 9 IV 1992, B. L, (CNC) (19); 16 III 1998, L. R, (CDRS) (1¢). Los Agua- cates, 16 III 1998, L. R (CDRS) (1d). NW slope 600 m, 30 V 1964, D. Q. Cavagnaro, (CAS) (1°).

Vanessa carye Hiibner [1812] “Western Painted Lady” (Figs. 15, 16) Hamadryas carye Hiibner {1812):pl. 45, figs. 1, 2. Pyrameis caryae; Williams 1911:300. Vanessa caryae; Linsley & Usinger 1966:156.

Diagnosis. FW = 21-28 mm. Differs from V. vir- giniensis by having four small blue-centered ocular markings on the VHW.

Geographical distribution. This species has been collected on San Cristdbal, Fernandina, and recently observed by the author in Floreana. Vanessa carye is distributed in the Andes of Colombia, Ecuador, Peru, Bolivia, Chile, southwestern Brazil, Paraguay, Uruguay, and Argentina. It also occurs on Juan Fernandez island, on Easter island, and the Tuamotu Archipelago.

Natural history. Three out of four specimens col- lected or observed were from the low arid zone at sea level. Two fresh adults were observed visiting flowers on Floreana (13 I 1995 and 6 IT 1995, Puerto Velazco Ibarra) but were not collected. Their fresh condition suggested that they were from a population extant on the island, rather than migrants. The host plant and the immature stages are unknown in the Archipelago.

Members of Asteraceae, Geraniaceae, Malvaceae, and Urticaceae are reported as host plants for the species (Field 1971).

Flowers visited. Verbenaceae: C. molle.

Flight period. Adults have been collected or ob- served in January and February.

Remarks. The only two specimens collected on the Galapagos were deposited in the California Academy of Sciences.

Material studied. (1¢ and 19). FERNANDINA: near to sum- mit, 4800 ft, 5 If 1964, D. Q. Cavagnaro, (CAS) (19). SAN CRISTO- BAL: Chatam, Wreck Bay, 22 IT 1906, (CAS) (1¢).

ACKNOWLEDGMENTS

I would like to thank the authorities of the Galapagos National Park and the Charles Darwin Research Station for financial and lo- gistical support in field. The Charles Darwin Research Station and Canacoll Foundation provided financial support for my investiga- tions at the Canadian National Collection of Insects in 1999. I thank the curators Germania Estevez (early curator MECN, Quito, Ecuador), Giovanni Onore and Francisco Pifias (PUCE), Bernard Landry (MHNG, Geneva, Switzerland) for their permission to study the specimens deposited in their collections. I am indebted to Bernard Landry and Charles V. Covell Jr. (University of Louisville, Kentucky, USA) for generosity with their time and knowledge of Lepidoptera, and Stewart Peck (Carleton University, Ottawa, Canada) for his knowledge of Galapagos insects. I am also greatly in- debted to Charlotte Causton, Alan Tye, Heidi Snell and Sol Rojas (CDRS), Olaf Mielke and Mima Casagrande (Universidad Federal de Parana, Parana, Brazil), Philip J. DeVries and Carla M. Penz (Milwaukee Public Museum, Milwaukee, USA) and Keith Willmott (NHM London) for their critical review of versions of this manu- script. To Ivan Aldaz (CDRS) for plant identifications, and Zsolt Balint (Hungarian Natural History Museum, Budapest, Hungary) for Lycaenidae literature. To Matts Wedin (CDRS) for producing the plates. Finally I thank to all my field assistants during the years, especially Valentina Cruz. i

This research was partially supported by Unesco project #894976.9: Ecological monitoring in the Galapagos Archipelago, Galapagos Conservation Fund, GEF-Natura Project: Monitoring the Galapagos Islands and Darwin Initiative for the survival of species project: Terrestrial Invertebrate Biodiversity in Galapagos (Ecuador): Training and collection rehabilitation.

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MCMULLEN, C. K. & D. M. VIDERMAN. 1994. Comparative studies on the pollination biology of Darwiniothamnus tenuifolius (Asteraceae) and Plumbago scandens (Plumbaginaceae) on Pinta Island and Santa Cruz Island, Galapagos. Phytologia 76 (1):30-38.

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Received for publication 19 June 2002; revised and accepted 7 Jan- uary 2004

Journal of the Lepidopterists’ Society 58(1), 2004, 44-47

HOST BREADTH AND VOLTINISM IN GALL-INDUCING LEPIDOPTERA

WILLIAM E. MILLER Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, USA, email: mille014@umn.edu

ABSTRACT. Because of specialized life systems and host relations, gall-inducing insects are believed to have narrower host breadth and lesser voltinism than other endophagous insects. These expectations were tested here using a sample of 136 species of British Lepidoptera con- sisting of 29 gallers in 11 families and 107 taxonomically matched endophagous nongaller controls. Gallers and nongallers were compared using biological data assembled and published by A. M. Emmet. Host breadth, scored as number of host genera, averaged 1.21 for the gallers, and 1.48 for the nongallers, and the difference is statistically significant, thus confirming host breadth expectation for this sample of Lepidoptera. This difference was generated mostly among Gelechiidae, Nepticulidae, Tortricidae, and Sesiidae. Voltinism, scored as number of generations developing per year, did not differ between ‘gallers and nongallers. This outcome does not necessarily refute the hypothesis underlying the ex- pectation. The cool British climate may inhibit multivoltinism, and thereby minimize differences in voltinism between gallers and nongallers.

Additional key words: Gelechiidae, Nepticulidae, Tortricidae, Sesiidae, Britain.

alls induced by Lepidoptera vary from simple See to fruitlike bodies that bear little resem- blance to supporting host parts. The galls are organ specific, most developing on host stems, but some de- veloping on host reproductive structures, leaves, and roots. With few exceptions, larvae rather than oviposit- ing adults induce the galls. Worldwide, 352 morpho-

species of gall-inducing Lepidoptera are known, of

which 179 have been identified to genera and species in 20 families (Miller 2004). Greater numbers and tax- onomic diversity of lepidopteran gallers are antici- pated as tropical areas are explored.

Gall-inducing insects, including Lepidoptera, are believed to have narrower host breadth and to develop fewer annual generations than other endophagous in- sects (Cornell 1990, Raman 1994, Miller 2004). Nar- row host breadth is expected because evolutionary ad- justments between gall inducer and host foster foodplant soaeiliantion, a corollary of the reputed host specificity of gall inducers (Mani 1964, Short- house & Rohfritsch 1992, Harris & Shorthouse 1996). Lesser voltinism is expected because gall inducers seem to synchronize their phenology with that of their hosts, which would ensure that larvae have access to reactive tissues necessary for gall development, as dur- ing rapid plant growth in spring. Physiological mech- anisms that might mediate this synchrony have not been inv estigated. These host breadth and voltinism expectations for lepidopteran gall inducers would ac- quire added strength if empirical tests confirmed them.

Cornell (1990) compared voltinism and other life history traits between gall inducers and leaf miners.

His voltinism sample consisted of 28 species—12 leaf

miners, mostly lepidopteran, and 16 gall inducers, mostly dipteran and hymenopteran. He found that vol- tinism averaged 1.4 generations/yr for the gall inducers and 2.5 generations/yr for the leaf miners, which is consistent with the expectation of lesser voltinism among gall inducers. In contrast to voltinism, host

breadth of gall inducers in one or any combination of insect orders does not seem to have been compared empirically with that of endophagous nongallers.

Reported here are comparisons of host breadth and voltinism between gallers and nongallers in a large sample of Lepidoptera, an order poorly represented in previous cecidological studies (Miller 2004). The source of the data analyzed is Emmet’s (1991) exten- sive life history tabulation for more than 2400 species of British Lepidoptera, the most extensively known lepidopteran fauna in the world.

MATERIALS AND METHODS

To test the hypotheses that gall-inducing Lepi- doptera have narrower host breadth and lesser voltin- ism than other endophagous Lepidoptera, I assembled a study sample of 29 gallers and 107 endophagous nongaller controls, 136 species in all. All known British

gallers were included, as listed by Spooner and Bow- a ey (1995), with emendations as follows: Argyresthia retinella Zeller, unaccountably absent from the list, was added (Robbins 1992), and Paranthrene tabaniformis rhinglaeforme (Hiibner), now considered synonymous with P. tabaniformis (Rottemburg), was removed (Spatenka et al. 1999).

The 107 endophagous nongallers were those marked in Emmet’s (1991) tabulation exclusively with b for borer or m for miner, and, for Heliozela only, also with c for casebearer to match the casebearing Heliozela gallers. Taxonomic matching was possible at the generic level for gallers in 10 genera, and at the sub- family level for gallers in six genera, in line with prin- ciples of the comparative method (Harvey & Pagel 1991). For example, matches for the two Boraad oat (Nepticulidae) gallers consisted of the 15 nongalling endophagous Ectoedemia, and the match for the galler Adaina microdactyla (Hiibner) (Pterophoridae, Platyptiliinae), which has no British congeners, was Leioptilus carphodactyla (Hiibner), the only other en- dophagous British member of the subfamily Platyptili-

VOLUME 58, NUMBER 1

TaBLE 1. Host breadth and voltinism of British gall-inducing Lepidoptera and taxonomically matched endophagous nongallers. Data from Emmet (1991) except where noted otherwise. Family sequence follows Kristensen (1999).

Mean scores

Family Species as numbered in Emmet’s tabulation N Host breadth Voltinism Nepticulidae Gallers 23, 24 2 1.00 1.00 Nongallers 25-32, 34-39, 41 15 1,27 1.00 Heliozelidae Gallers 154, 157 2) 1.00 1.00 Nongaller 156 1 1.00 1.00 Incurvariidae Gallers 138, 139 2 1.00 1.00 Nongallers 133, 136 2 1.00 1.00. Yponomeutidae Gallers 411, 415 2 1.50 1.00 Nongallers 401, 404, 405, 407, 410, 412, 418, 420, 422 1e33 1.00 Elachistidae Galler 906 1 1.00 00 Nongaller 905 1 1.00 1.00 Coleophoridae Gallers 486, 889, 891, 892, 893a! 1.00 1.25 Nongallers 487, 880-884, 887, 888, 890 9 1.00 33 Gelechiidae Gallers 728, 755 2 1.00 1.00 Nongallers 723-727, 727a, 729, 730, 735, 737, 744, 744a, Q7 wall 138} 746-748, 753, 757, 808, 811-813, 816, 817, 821, 822, 823a, 825 Sesiidae Gallers 372, 377, 380 3 1883 0.67 Nongallers 373-379, 381 8 1.50 0.75 Tortricidae Gallers 966, 1137, 1167, 1190, 1195, 1256, 1258, 1266 8 Wit 1.06 Nongallers 962, 964, 965, 967, 1168, 1192, 1194, 1196, 1197, 34 1.56 1.06 1199, 1200, 1200a, 1201, 1202, 1240, 1242, 1243, 1245-1247, 1249, 1253-1255, 1257, 1259-1261, 1264, 1265, 1267, 1268-1270 Pterophoridae Galler 1517 i 1.00 2.00 Nongaller 1519 ik 2.00 2.00 Crambidae Galler 1359 1 2.00 1.00 Nongallers 1375 1 1.00 2.00 Summary Gallers 29 1.21 1.07 Nongallers 107 1.48* 1.11

‘Mompha bradleyi Riedl, whose discovery in Britain (Harper 1994) postdates Emmet (1991).

* Mann-Whitney U ,,,3-= 1280.0, Pp, saijeq < 0-05.

134df

inae. Nongallers outnumber gallers in the study be- cause plausible matches were often more numerous than the gallers matched, all being included to avoid selection bias.

Host breadth and voltinism data were extracted for both the gallers and nongallers from Emmet’s (1991) tabulation. Data for one galler subsequently discov- ered in Britain, Mompha bradleyi Riedl, was obtained from Harper (1994). Host breadth was scored as num- ber of recorded host genera. This is a stringent mea- sure in that no distinction was made between one and

more than one host species in the same genus; how- ever, the problem of appropriately scaling and inte- grating genus and species scoring was thereby avoided. Scoring by species alone could not be done because the source did not consistently list numbers of host species within genera. Voltinism was scored as number of annual generations, with the case of less than one annual generation (one generation every two years) being scored as 0.5. This case had minimal impact be- cause it occurred in only 3 of the 29 gallers (2 sesiids and | tortricid) and 4 of the 107 nongallers ( all sesiids).

46

Student’s t, Mann-Whitney U, and statistical sum- maries were computed with SYSTAT (1992) software. Homogeneity of variance between galler and nongaller groups was examined before analysis as outlined by

Sokal and Rohlf (1981).

RESULTS

The 29 British gall inducers represent 11 of the 20 families of identified lepidopteran gall inducers world- wide (Table 1; Miller 2004). Stem galls are induced by 21 of the gallers, petiole galls by 5, and reproductive- structure galls by 3 (Robbins 1992, Spooner & Bow- drey 1995).

Mean host breadth was 1.21 genera for gallers com- pared to 1.48 genera for endophagous nongallers. The difference, 0.27, is in the expected direction of fewer host genera for gall inducers, and is significant (U-test, Table 1). The nonparametric U-test was used because galler and control variances proved divergent. Host breadth ranged 1-2 for the gallers, and 1-4 for the nongallers.

The difference in host breadth between the two groups originated mainly within Gelechiidae, Nepticul- idae, Tortricidae, and Sesiidae. In these families, host breadth means for nongallers exceeded those for gallers by 0.71, 0.27, 0.19, and 0.17, respectively (Table 1).

Mean voltinism was 1.07 annual generations for the gall inducers, compared to 1.11 annual generations for the nongallers (Table 1). The difference, 0.04, al- though in the expected direction, is not significant (t = —0.53, P.. saiteq = 0.30). Galler and nongaller variances for voltinism proved to be homogeneous, which per- mitted the use of the parametric t-test. Voltinism ranged 0.52 for both gallers and nongallers.

DISCUSSION

Analysis of number of host genera supports the ex- pectation of narrower host breadth for gall inducers than for endophagous nongallers among British Lepi- doptera. This result (Table 1) might have been stronger had more elaborate scoring captured the intrageneric component of host breadth, but no objective method for combining or integrating genus and species scoring was available, and species counts alone could not be used because the source did not fully elaborate host species. Actual host breadth might also be stronger than indicated by the analysis for yet another reason: galls undoubtedly attract more interest and attention than other signs of insect infestation, so that host breadth of nongallers might be underreported compared to that of gallers, which could reduce the apparent difference.

Narrow host breadth is a trait that is highly desir- able in biological control agents of weeds, one which,

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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

other things being equal, enhances the biological con- trol potential of gallers (Harris & Shorthouse 1996, McEvoy 1996). Gelechiidae and Tortricidae, which are among the four families here with strongest differ- ences between gallers and nongallers in host breadth (Table 1), are also the families with the greatest num- bers of known gallers (Miller 2004).

Analysis does not support the expectation of lesser voltinism among gallers. This result (Table 1) contra- dicts Cornell's (1990) finding of lesser voltinism in gallers than leaf miners, but does not necessarily re- fute the hypothesis giving rise to the expectation. The British climate is cool-temperate and rainy, with the warmest month averaging less than 22°C (Lewis 1994). Such a climate may inhibit multivoltinism so that dif- ferences in voltinism between gallers and nongallers are minimized. By contrast, in eastern North America, a temperate land mass much larger than Britain, lepi- dopteran voltinism markedly increases as latitude de- creases (Fracker 1920, Tauber et al. 1986). Multivol- tinism in lepidopteran gall inducers, when it occurs, seems influenced more indirectly than directly by cli- mate; that is, protracted g growth seasons in warm tem- perate regions and in tropical wet cycles expand tem- poral av: ailability of reactive host tissues for gall development (Miller 2004). Although Cornell (1990) did not taxonomically match gallers and leaf miners in comparing their voltinism, he reported no statistical connection between taxonomy and analytical outcome, and his result unequivocally shows lesser galler voltin- ism. Further testing of the voltinism expectation would be desirable. Possible mechanisms underlying galler— host phenological synchrony might include higher thermal thresholds for galler than nongaller develop- ment, and differing diapause reactions.

ACKNOWLEDGMENTS

I thank R. C. Venette, D. W. Ragsdale, R. W. Hodges and K. A. Schick for useful reviews of the manuscript.

LITERATURE CITED

CorNELL, H. V. 1990. Survivorship, life history, and concealment: a comparison of leaf miners and gall formers. Amer. Nat. 136: 581-597.

EmMe_T, A. M. 1991. Chart showing the life history and habits of the British Lepidoptera, pp. 61-301. In Emmet, A. M. & J. Heath (eds.), The moths and butterflies of Great Britian and Ireland. Vol. 7, Pt 2. Harley, Colchester, United Kingdom.

FRACKER, S. B. 1920. The life cycle of the Lepidoptera. Ann. En- tomol. Soc. Amer. 13:167-173.

Harper, M. W. 1994. Mompha bradleyi Riedl (Lepidoptera: Mom- phidae) new to Britain, with some initial observations on its life history. Entomol. Gaz. 45:151-156.

Harris, P. & J. D. SHoRTHOUSE. 1996. Effectiveness of gall induc- ers in weed biological control. Can. Entomol. 128:1021—-1055.

Harvey, P. H. & M. D. PAGEL. 1991. The comparative method in evolutionary biology. Oxford, Oxford, United Kingdom.

VOLUME 58, NUMBER 1

KRISTENSEN, N. P. (ED.). 1999. Lepidoptera, moths and butterflies. Evolution, systematics, and biogeography. Handbook of Zool- ogy. Vol. 1. De Gruyter, New York.

Lewis, H. A. G. 1994. The Times atlas of the world. 9th ed. Times Books, London.

Mani, M. S. 1964. Ecology of plant galls. Dr Walter Junk, The Hague, The Netherlands.

McEvoy, P. B. 1996. Host specificity and biological control. BioSci. 46:401405.

MILLER, W. E. 2004. Gall-inducing Lepidoptera, pp. 429-462. In Raman, A., C. W. Schaefer & T. M. Withers (eds.), Biology, ecology, and evolution of gall-inducing arthropods. Science Publishers, Enfield, New Hampshire, USA.

Raman, A. 1994. Adaptational integration between gall-inducing insects and their host plants, pp. 249-275. In Ananthakrishnan, T_N. (ed.), Functional dynamics of phytophagous insects. Sci- ence Publishers, Lebanon, New Hampshire, USA.

RosBINs, J. 1992. Argyresthia retinella Zell. (Lepidoptera: Ypono- meutidae) a gall causer. Cecidology 7:53.

SHORTHOUSE, J. D. & O. ROHFRITSCH (EDS.). 1992. Biology of insect- induced galls. Oxford University Press, New York.

SOKAL, R. R. & F. J. ROHLF. 1981. Biometry. 2nd ed. Freeman, New York, USA.

SPATENKA, K., O. GORBUNOY, Z. LASTUVKA, I. TOSEVSKI & Y. ARITA. 1999. Handbook of Palaearctic macrolepidoptera. Vol. 1. Sesi- idae—Clearwing moths. Gem, Wallingford, England.

SPOONER, B. M. & J. P. Bowdrey. 1995. Checklist of British galls and gall-causing organisms. 1. Lepidoptera: preliminary list. Cecidology 10:84-100.

SYSTAT. 1992. Statistics version 5.2 ed. SYSTAT Inc.

TAuBER, M. J., C. A. TAUBER & S. MASAKI. 1986. Seasonal adapta- tions of insects. Oxford, New York.

Received for publication 10 March 2003, revised and accepted for publication 19 September 2003

GENERAL NOTES

Journal of the Lepidopterists’ Society 58(1), 2004, 48-50

NEW RANGE EXTENSIONS, LARVAL HOSTPLANT RECORDS AND NATURAL HISTORY OBSERVATIONS OF CUBAN BUTTERFLIES

Additional key words: Euphorbiaceae, Fabaceae, Bignoniaceae, Acanthaceae, Rutaceae.

Although butterflies have not been thoroughly col- lected in Camagiiey province, a checklist of the terri- tory has been recently published as the result of field surveys and collections completed during the last ten years (Fernandez & Rodriguez 1998). The latter com- plements the preliminary work of Guerra et al. (1993), who reported 76 butterflies and skippers. This up- dated and augmented list included 125 species, but their residency status has not been entirely deter- mined within the province due to the paucity of infor- mation available for some of them. A few have only been collected or observed on one or two occasions due to the short collecting time available.

Here, I report range extensions for the nymphalid Antillea pelops anacaona (Herrich-Schiffer), the ly- caenid Strymon acis casasi (Comstock & Huntington) and the hesperiid Achlyodes munroei Bell, which rep- resent new butterflies for the province. The associated larval hostplants for these species and for six other but- terflies and biological, ecological and distributional

data are also discussed. Voucher adult specimens of

the three newly recorded species are deposited in the author's collection and a pair of A. munroei with asso- ciated pupal exuvia has been deposited in the collec- tion of the Allyn Museum of Entomology, Florida Mu- seum of Natural History.

These observations were made between May 1996 and November 2001 in the following localities in Cam- agiiey province: Albaiza, Camagiiey city, Central Can- dido Gonzalez, Limones, Loma de Yucatan and Santa Ana (south of the INIVIT Experimental Station). The fourth and the fifth sites were areas not previously sampled. Additional information was also obtained from other field trips at the southern slope of Sierra de Cubitas in 1988, 1994 and 1996.

NYMPHALIDAE

Anaea cubana (H. H. Druce). One fifth and four fourth instar larvae were found in the leaves of Croton sagraeanus Muell. Arg. (Euphorbiaceae) at Limones in June. All were inside vertical tubular shelters, open at both ends, and formed by tying together various longitudinally interconnected leaves. The caterpillar always enters the upper and wider opening and backs into the tube (caudal end first) once it returns from

feeding. Once inside, the conspicuous head capsule remains exposed at the entrance. The fourth instar lar- vae were fed with the leaves of Croton argenteus L. in- stead of the original host. They accepted the new food- plant and completed development to the adult stage. On this plant, the larvae either lived’ exposed or tied the edges of an individual leaf for shelter. Croton ar- genteus was growing spontaneously along roadsides in the surroundings of the city of Camagiiey. Pupation took place in the underside of the leaves of the host (in the field, a pupal exuvia was also found on the under- side of a C. sagraeanus leaf) and lasted 9 (n = 2 males) and 10 (n = 2 females) days. The adults were not abun- dant at the collecting site but seen during all of the field visits, flying along hedgerows and roadsides and attracted to fallen, ripe mangoes.

According to Alayo an Hernandez (1987), the larva of A. cubana was reared by Gundlach on Pectis (Asteraceae) species. As all other species of Anaea have been reported to be confined to Euphorbiaceae (Smith et al. 1994), this record on Asteraceae is likely a mistake.

Hamadryas amphichloe diasia (Fruhstorfer). Two larvae were found on leaves of Dalechampia scan- dens L. (Euphorbiaceae) at Loma de Yucatan. There were not records of foodplants for this Antillean sub- species but Smith et al. (1994) mentioned that species of Dalechampia or Tragia were likely to serve as host.

Antillea pelops anacaona (Herrich-Schiffer). Seven specimens of this endemic subspecies (five males and two females) were taken at Limones in June and September. These were found in humid, shady undergrowth beneath a mango tree in a wooded area. The individuals were perching on plants less than 1 m tall, with both wings slightly, partly or fully open (but not pressed against t the leaves). Sometimes the butter- flies flexed their wings at a rapid rate or were seen fly- ing just above or between the low vegetation. At times, the flight may be slow and weak, but it can also be quite agile and difficult to follow. All these adult be- haviors combined to make the butterfly inconspicuous supporting the observations made by Smith et al. (1994) that they may be overlooked unless present in numbers. A male nectaring on Alternanthera axillaris (Hornem.) D. Dietr. (Amaranthaceae) and a specimen

VOLUME 58, NUMBER 1

thermoregulating with wings partly open before mid- day were other hehavions observed for the butterfly. Two fourth instar larvae were independently col- lected on Blechum pyramidatum (Lam.) Urb. and Jus- ticia comata (L.) Lam. (both Acanthaceae) in Septem- ber. One of the larvae pupated and emerged as an adult male after six days. The butterfly appears to be resi- dent, and on the last two visits to the area in Septem- ber, I observed two fresh pairs. This tiny nymphalid was reported by Alayo & Hernandez (1987) in Gran Piedra and Sardinero, Santiago de Cuba, in eastern Cuba: and Pinares de Vinales and Luis Lazo, Pinar del Rio, to the west. They are always seen in small isolated groups. Fontenla (1987) also reported it from a wooded area of Parque Lenin, Ciudad de La Habana.

LYCAENIDAE

Strymon acis casasi (Comstock & Huntington). Three adults and a last instar larva of this endemic subspecies were taken at Limones in May and June. The first specimen found was a rather worn female that perched on a low-growing plant of Dichrostachys cinerea Willd. (Fabaceae). It was collected just after an erratic and instantaneous flight along a path between dense groups of the exotic ‘Vege, The two other specimens, a badly worn female which was released and a fresh male, were collected on C. sagraeanus flowers. A larva was also collected in the same area as the adults and was reared on the Croton flowers until pupation. However, the adult did not emerge. Other lycaenids, including many male Ministrymon azia (He- witson) and some Strymon columella cybira (Hewit- son), Leptotes cassius theonus (Lucas), Hemiargus hanno filenus (Poey) and Cyclargus a. ammon (Lucas) were also present and nectaring on the Croton flowers. S. acis casasi was not seen commonly, and more visits to the locality will be required for further observations to determine the peak months of adult activities. Alayo and Hernandez (1987) cited this subspecies as rare, being only collected around Guantanamo province or some other localities of the extreme eastern part of the island.

Strymon columella cybira (Hewitson). A last in- star larva was found on C. argenteus flowers in Cam- agiiey city. In Cuba, a wide range of foodplants is now known for this lycaenid, not only in the Malvaceae but also members in the Acanthaceae, Portulacaceae, Ster- culiaceae and Surianaceae (Fernandez 2001).

Leptotes cassius theonus (Lucas). Larvae were al- ways found feeding on flowers. In Camagiiey city, lar- vae were on Albizzia lebbeck (L.) Benth., Calliandra surinamensis Benth. and Pithecellobium dulce Benth. (Fabaceae). In Albaiza, larvae were on Calopogonium mucunoides Desy. (Fabaceae), and in Central Candido

49

Gonzalez, they were on Clytostoma callistegioides Bur. (Bignoniaceae), a plant family not previously recorded for the lycaenid. The hostplants for L. cassius are gen- erally herbaceous legumes but Malpighiaceae and Plumbaginaceae are als used (Smith et al. 1994).

HESPERIIDAE

Proteides maysi (Lucas). Oviposition on the un- derside of a leaflet of Andira inermis (Wright) Kunth ex DC. (Fabaceae) was observed at Limones. After eclo- sion, the larva was reared on Lonchocarpus dominguen- sis (Turp. ex Pers.) DC. (Fabaceae) through the fifth in- star but died due to excessive humidity in the rearing container. No previous life history information was avail- able for this endemic Cuban skipper (Smith et al. 1994).

Burca concolor concolor (Herrich-Schiffer). The larvae of different instars were commonly ob- served on C. sagraeanus leaves at Limones in May, June and September. They construct shelters by using interconnecting leaves or bending them inward and sealing the unions between margins, and using a to- mentum present on the underside of leaves. Earlier in- star larvae build a shelter on the ventral surface of a single leaf just using this tomentum and feed on leaf edges. Active feeding occurs only on the upper side (leaving the tamara) which produces the effect of a chlorotic or scraped leaf. Later instar larvae cut out the leaf section where they have fed so damaged parts are not visible on the plant. Pupation takes ‘place in the shelter and lasts 8-9 days (n = 3).

The adults were more numerous only in June. The males constantly patrolled around and between Croton stands at low or medium height from the ground. The females also joined males in this behavior with oviposi- tions in some cases; one female was also taken while perching with closed wings on a Gliricidia sepium (Jacq.) Kunth ex Walp. (Fabaceae) leaflet to a height of about 4 m. Individuals of both sexes were seen taking nectar with wings fully open on flowers of the hostplant, but males also visited Koanophyllon villosum (Sw.) R.M. King et H. Rub. (Asteraceae). Solitary males were observed sipping water at the edges of mud puddles. This skipper appears to be resident and fairly common at Limones.

In addition, several larvae were also found on Cro- ton origanifolius Lam. at Albaiza in August and No- vember. Here, no adults of this species had been recorded during previous field studies but upon locat- ing the immatures, a female was observed in the nearby hostplant patch at a latter date. This individual took nectar on C. origanifolius and on Bourreria mi- crophylla Griseb. (Boraginaceae), and later alighted with wings open on dry grasses on the ground, where its presence was inconspicuous.

50.

Burca c. concolor was recorded for the first time in Camagiiey in 1988 near Paso de los Paredones in the dry plains of the southern slope of Sierra de Cubitas hills where the species was seen along roadsides. In more recent years (1994 and 1996), it has also been found there again and seen at times in abundance. In- dividual specimens have been observed alighting on oe or low plants and nectaring on Waltheria indica

L. (Sterculiaceae) flowers. In Rddigon to the above mentioned localities, other specimens of this skipper have been also taken in a wooded tract around the lower part of Loma de Yucatan in June. Due to its general as- sociations with grasses and open areas, this species is probably originating from the adjacent savannas.

This endemic subspecies is considered as very rare in Cuba (Alayo & Hernandez 1987), recorded from pico Potrerillo, Escambray; Cuabales de Corral Nuevo, Matanzas; Tortuguilla, Guantanamo; playa Juragua, Santiago de Cuba. Smith et al. (1994) have added playa Ameen near Trinidad as another collecting site. The

discovery of the foodplant association ath Croton of

this species will not only help to understand the signif- icance of the local occurrence of the skipper but may perhaps encourage a wider search of this species in lo- cal foodplant patches.

Achlyodes munroei Bell. Miller and Simon (1998) mentioned that apparently the last known spec- imens of A. munroei were collected over a half century ago by Pastor Alayo, despite continued diligent search for it in Cuba by Luis Roberto Hernandez and others. Some lamented that this insect might be extinct. Al- though older A. munroei specimens were found in the coastal locality of Siboney, Santiago de Cuba (Smith et al. 1994), the area for the recently collected ones is an inland site, primarily a cattle pasture with many trees and low-growing bushes, including two Zanthoxylum. Here A. mithridates papinianus (Poey) is amore com- mon inhabitant and found in various larval stages, which utilize all rutaceous trees available (including Citrus). This area remains abandoned for some peri- ods that allows plant regrowth and formation of very suitable habitats, not only for resident butterflies but for temporary colonizers, such as A. munroei.

I found this species as larvae in leaf shelters of Zan- thoxylum martinicense (Lam.) DC. (Rutaceae) in Au- gust at Santa Ana. A male was captured in November

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

2001, at the same site. Apart from the perching pos- ture of the third collected specimen, with wings fully open and resting on leaves at about 40 cm high from the ground, no other distinct behaviors were seen.

Until very recently, A. munroei was considered to be an endemic Cuban species, but it has subsequently been found on Cat Island, Bahamas. Recent collec- tions there suggest that the insect is well established and will probably be recorded on other Great Bahama Bank islands in the future (Miller & Simon 1998). The discovery of the species in Camagiiey not only con- firms that A. munroei still exists in Cuba, but it may well prove to be also more widely distributed as fur- ther field studies are conducted by resident entomolo- gists.

I wish to thank Adelaida Barreto (CIMAC, Camagiiey) and Eddy Martinez (BIOECO, Santiago de Cuba) for their invaluable assis- tance in the identifications of the larval foodplants and nectar sources. Special thanks to Drs. Lee and Jacqueline Miller (Allyn Museum of Entomology) for confirming identification of A. munroei, helping with the English and reviewing the manuscript. My wife Noris V. Verdecia and Carlos A. Sacasas also helped during preparation of the manuscript and warrant my thanks.

LITERATURE CITED

Aayo, P. & L. R. HERNANDEZ. 1987. Atlas de las mariposas diur- nas de Cuba (Lepidoptera: Rhopalocera). La Habana, Ed. Cientifico-Técnica. 148 pp.

FERNANDEZ, D. M. 2001. New oviposition and larval hostplant records for twenty-three Cuban butterflies, with observations on hee biology and distribution of some species. Carib. J. Sci. 37(1-2):122-125.

ait nines D. M. & L. RopricuEz. 1998. Las mariposas de Ca- magiiey (Lepidoptera: Papilionoidea y Hesperioidea). Cocuyo 7:21-23.

FONTENLA, J. L. 1987. Aspectos comparativos estructurales de tres comunidades de mariposas (Lepidoptera: Rhopalocera) en Cuba. Poeyana 337:1-20.

Guerra, M., R. RODRIGUEZ, E. VELAZQUEZ, R. ACOSTA, E. SUAREZ & H. Sonora. 1993. Listado preliminar de las mariposas diur- nas (Lepidoptera: Rhopalocera) de Camagiiey. Monteverdia 1(1):10-13.

MILLER, L. D. & M. J. SIMON. 1998. Rediscovery of the rare, “en- demic” Cuban butterfly, Achlyodes munroei, with notes on its behavior and possible origin. Carib. J. Sci. 34(3-4):327-329.

Smitu, D. S., L. D, MILLER & J. Y. MILLER. 1994. The butterflies of the West Indies and South Florida. New York, Oxford Univ. Press. 264 pp.

DouGLas MANUEL FERNANDEZ HERNANDEZ, Apartado Postal 83, Camagiiey 1, C.P. 70100, Cuba, email: douglasm@medscape.com

Received for publication 16 December 2002, revised and accepted 17 September 2003.

Journal of the Lepidopterists’ Society 58(1), 2004, 51-53

IMMATURE STAGES OF CALYDNA VENUSTA MORIO (RIODINIDAE) FROM TRINIDAD

Additional key words: _ balloon setae, Olacaceae, Ximenia americana.

Trinidad, which forms part of Trinidad and Tobago, has a long history of entomological exploration, and successive generations of resident and visiting collec- tors have together compiled a reasonably complete picture of the butterfly fauna of this species-rich, con- tinental island. The landmark publication for the coun- try, Barcant’s (1970) “Butterflies of Trinidad and To- bago”, recognized 103 riodinid species from Trinidad. Nevertheless, new records of Riodinidae continue to be added, and the authors are preparing an updated checklist. One of these additions is Calydna venusta morio Stichel (Riodininae: incertae sedis section), dis- covered in 1984 by S. Alston-Smith at Point Gourde, on the northwestern peninsula of Trinidad, as indi- cated in D’Abrera (1994) and Hall