PROC. ENTOMOL. SOC. WASH. 90(2), 1988, pp. 164-178

DESCRIPTION AND BIOLOGY OF ACROLOPHUS PHOLETER, (: TINEIDAE), A NEW COMMENSAL FROM GOPHER TORTOISE BURROWS IN FLORIDA

(DRD) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (EGM) U.S. Army Environmental Hygiene Agency, Field Support Activity, Fort Mead, Maryland 20755.

Abstract. -Vacuum sampling of gopher tortoise burrows in Putnam County, Florida, has resulted in the discovery of a new species of tineid moth, Acrolophus pholeter Davis. The larva feeds on both the fecal pellets of the gopher tortoise and upon decaying plant debris within the burrow. Supplemented by numerous illustrations, the larval, pupal, and adult stages are described, and the general biology is summarized. Key Words: Lepidoptera, Tineidae, moth biology, Gopherus polyphemus, gopher tor- toise

Recent vacuum sampling for invertebrate this tortoise or if the moth also frequents commensals in the burrows of the gopher rodent burrows. The absence of previous tortoise, Gopherus polyphemus Daudin, by Acrolophus collecting records suggests that the junior author has revealed the presence its habitat may be rather restricted. of a few previously unreported. Since at least the Pleistocene, gopher tor- Among these was a new species of Acrolo- toise burrows have provided a relatively phus that was found in abundance feeding stable habitat for the establishment of a di- on both tortoise fecal pellets and decaying verse community of organisms. The integ- plant debris within the burrow. rity of individual burrows is normally main- This is the first record of an Acrolophus tained for five years or more. In terms of commensal in an burrow. The sub- numbers of both vertebrate and inverte- terranean, tube-constructing habit of the ge- brate species found using gopher tortoise nus, however, is well known. Other Tine- burrows, the diversity is one of the greatest idae (all Acrolophinae) have been reported yet studied in North American animal bur- from rodent burrows (Hubbard 190 1, Hub- rows (Milstrey 1986). bell and Goff 1939, Davis et al. 1986), and Sampling of organisms from gopher tor- at least one other moth, Idia gopheri (Smith), toise burrows can be a formidable task. De- is known to inhabit the burrows of the go- pending upon soil type and water table, bur- pher tortoise (Hubbard 1894, 1896, Smith rows may extend up to 40 feet long and 12 1899, Woodruff 1982). Hubbell and Goff feet deep (Young and Goff 1939). Excava- (1939) reported that some com- tion of such galleries can create a sizeable mensals were true obligates and had not been trench (Hubbard 1894). In recent years the collected outside gopher tortoise burrows. use of vacuum suction devices (Butler et al. It is not known to what extent Acrolophus 1984) has greatly facilitated collecting from pholeter n. sp. is restricted to burrows of burrows without decimating the landscape VOLUME 90, NUMBER 2 165

Figs. 1-5. Acrolophus pholeter, habitat, adults, and cocoon. 1, Entrance to gopher tortoise burrow, Gopherus polyphemus, Putnam Co., Florida. 2, Modified Echo R200 blower used for sampling invertebrate fauna from burrows. 3, Adult male, length of forewing 6.7 mm. 4, Adult female, length of forewing 8.2 mm. 5, Cocoon with pupal exuvium protruding, length of cocoon 13 mm. 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Fig. 2). One disadvantage of this technique uniformly brownish gray, relatively smooth is that it does not allow direct observations vestiture with slightly rough scales along of the organisms' biology. venter of second segment. All material used in this study was col- Thorax: Pronoturn and both fore and lected by the junior author using a modified hindwings uniformly brownish gray. Venter Echo R200 (R) blower (Kioritz Corp., To- somewhat paler, more light brown. Legs with kyo, Japan). Most adults (all type material) smooth vestiture, light brownish gray dor- were reared from larvae collected by vac- sally, lighter buff ventrally; epiphysis greatly uuming. reduced (Fig. 14), length only about twice Deposition of specimens referred to in its width. Pretarsus of all legs unspecialized, this paper are: BMNH for British Museum with symmetrical claws, pulvilli, well de- ofNatural History, London, England; FSCA, veloped arolium, and unguitractor plate Florida State Collection of Arthropods, bearing 5-6 transverse rows of scutes. Gainesville, Florida; and USNM, National Abdomen: Uniformly light brownish gray. Museum of Natural History (formerly Male genitalia: As shown in Figs. 18-21. United States National Museum), Smith- Uncus elongate, slender, and acute. Tegu- sonian Institution, Washington, D.C. men relatively broad and elongate. Vincu- lum slender; anterior margin slightly con- Acrolophus pholeter Davis, cave at middle. Gnathos a well developed NEW SPECIES median lobe with a relatively broad, trun- Figs. 3-62 cate apex. Valva rather broad over basal Adult (Figs. 3-4). -Length of forewing: half to sharply defined saccular lobe, male, 5.5-6.7 mm; female, 7-9.5 mm. A abruptly narrowing beyond lobe to simple moderately small moth with uniformly apex. Aedoeagus relatively short, approxi- brownish gray wings, smooth head, and mately two-thirds the length of valva, and short labial palpi. without cornuti; apex with a serrated cleft Head: Vestiture smooth over vertex and extending over one-fourth down right side. frons; scales uniformly brownish gray, rel- Female genitalia: As shown in Fig. 22. atively slender with rounded apices, ap- Only a single pair of short, posterior apophy- pearing to arise from lower frons and curv- ses present. Caudal margin of lamella an- ing upwards over vertex until they reach tevaginalis smoothly curved. Ductus bursae flattened, transversely oriented scale patches very short, slightly thickened and constrict- across occiput. Eye small, interocular index ed just before corpus bursae; latter simple, approximately 0.6 5, cornea relatively relatively small, membranous sac without smooth with only scattered microsetae (Figs. spicules. 10-1 1); eyelash absent. Antenna about 0.4- Larva (Figs. 33-62). -Length of largest 0.6 the length of forewing, relatively longer larva 18 mm; diameter 2.1 mm. Body trans- in male, 46-49 segmented; scape uniformly lucent, light yellowish brown with light brownish fuscous, smooth, without pecten; brown thoracic and anal plates. flagellomeres of similar color, fully scaled, Head: Uniformly light reddish brown, simple in form with a few sensilla coelo- darker around base of mandibles. Greatest conica along anterior margin (Figs. 12-1 3). width 1.4 mm, length 0.9 mm. AF2 arising Pilifer reduced, minutely setose (Fig. 7). well above (caudad) apex of frons. P2 more Mandible absent. Maxillary palpus greatly distant from P1 than P1 is to ecdysial line. reduced, 2 segmented; approximately 8-10 Stemmata vestigial, probably non-function- elongate sensilla arising from apical pit. al; only three transparent vestiges remaining Haustellum absent. Labial palpus short; (Fig. 53); one situated above S2, a very small length approximately 2.5 x eye diameter, one well below S2, and an elongate hyaline VOLUME 90, NUMBER 2 167

Figs. 6-1 1. Adult structure, Acrolophuspholeter. 6, Partial view of frons and mouthparts (88 pm). 7, Maxillary palpi and pilifers (37.5 pm). 8, Sensilla at apex of maxillary palpus (2.5 pm). 9, Sensilla at apex of labial palpus (8.8 rtm). 10, Eye (88 pm). 11, Detail of cornea showing scattered interfacetal microsetae (19 pm). (Scale lengths m parentheses.) 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 12-17. Adult structure, Acrolophuspholeter. 12, Antenna near middle of flagellum (37.5 pm). 13, Detail of Fig. 12 showing sensilla coeloconica (16.5 pm). 14, Reduced epiphysis on foretibia (25 pm). 15, Detail of epiphysis (7.5 pm). 16, Pretarsus of hindleg (19 pm). 17, Detail of unguitractor plate (3.75 pm). (Scale lengths in parentheses.) VOLUME 90, NUMBER 2 169

Figs. 18-24. Acrolophuspholeter. 18, Male genitalia, ventral view (0.5 mm). 19, Lateral view. 20, Lateral view of valva. 21, Lateral view of aedoeagus. 22, Female genitalia, ventral view (0.5 mm). 23, Pupa, ventral view (2 mm). 24, Dorsal view. (Scale lengths in parentheses.) 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 25-30. Acrolophus pholeter; pupa. 25, Head, ventral view (0.3 mm). 26, Lateral view (0.22 mm). 27, Caudal end (A7-10) of abdomen (0.22 mm). 28, Lateral view, dorsum up (0.19 mm). 29, Dorsal view (0.22 mm). 30, Detail of dorsal cremaster, A10 (88 pm). (Scale lengths in parentheses.) VOLUME 90, NUMBER 2 171

Figs. 31-36. Acrolophus pholeter. 31, Pupa, dorsum of A4-5 (0.43 mm). 32, Detail of spine row, A5 (30 pm).33, Larva, dorsal view of head (0.19 mm). 34, Dorsal view of labrum and mouthparts (68 pm). 35, Lateral view of head (0.17 mm). 36, Lateral view of stemmatal area (60 km). (Scale lengths in parentheses.) 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 37-42. Acrolophus pholeter, larva. 37, Head, frontal view (0.15 mm). 38, Maxillae and labium (68 pm). 39, Detail of maxilla (22 pm). 40, Ventral view of maxillae and labium (88 pm). 41, Detail of spinneret and labial palpi (37.5 pm). 42, Detail of secondary labial setae in Fig. 4 1 (3.75 km). (Scale lengths in parentheses.) VOLUME 90, NUMBER 2 173

Figs. 4348. Acrolophus pholeter, larva. 43, Antenna (43 pm). 44, Detail of antenna1 apex (12 bm). 45, Lateral view of prothorax (0.22 mm). 46, Ventral view of prothorax (0.22 mm). 47, Tarsal claw (1 1.5 bm). 48, Prolegs, A5 (0.15 mm). (Scale lengths in parentheses.) 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 49-54. Acrolophuspholeter, larva. 49, Crochets on proleg 5 (38.5 rm). 50, Anal proleg, A10 (60 rm). 5 1, Segments A9-10, dorsal view (0.19 mm). 52, Lateral view (0.19 mm). 53, Ventral view (0.19 mm). 54, Caudal view (0.19 mm). (Scale lengths in parentheses.) VOLUME 90, NUMBER 2

Figs. 55-62. Acrolophuspholeter, larva. 55, Chaetotaxy ofbody, segments TI-2, Al, 6,8-9. 56, Head, dorsal view (0.5 mm). 57, Ventral view. 58, Segments A%-10, dorsal view. 59, Head, lateral view. 60, Labrum, dorsal view (0.2 mm). 61, Ventral view. 62, Mandible (0.2 mm). (Scale lengths in parentheses.) area anterior to S2; the latter apparently the Mandible somewhat tapered, with 4 small remnant of three fused stemmata. Apical cusps. Maxilla as in Figs. 38-39. Spinneret segment of antenna relatively long; sensilla elongate, slender; labial palpus 2-segment- as in Figs. 43-44. Labrum with M1 and 2 ed, basal segment elongate with a short api- near anterior margin; M3 more remote. cal seta; apical segment greatly reduced, 176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON about 0.25 the length of basal segment, with em. 11 May 1985; 2 8, 3 0, em. 12 May an elongate seta nearly equal in length to 1985; 4 a, 1 a, em. 17 May 1985; 1 a, 12 both segments; apex of mentum with a pair June 1985, em. 30 June 1985; 1 2, em. 20 of minute secondary labial setae (Figs. 4 1- July 1985; 1 a, 10 July 1985, em. 23 July 42). 1985; 60 larvae, 26 July 1985; 68 larvae, 9 Thorax: Pronotal and mesonotal plates Sept. 1985. Paratypes deposited in BMNH, light brown. Spiracular plate almost com- FSCA, and USNM. pletely separated from pronotal plate; all 3 Host.-Larval substrate consists of both lateral setae together on spiracular plate. decaying plant debris within burrow of go- Coxal plates separated slightly. Tarsal claw pher tortoise and fecal pellets of tortoise. as in Fig. 47. Flight period. - Difficult to assess; adults Abdomen: A1-6 with 11 pairs of primary were collected from burrows through much setae, SV trisetose. Ventral crochets in a of the year. uniserial ellipse of approximately 30 hooks; Distribution. -Known only from under- sides of proleg densely covered with small, ground burrows of the gopher tortoise in the scattered spines, anal proleg with approxi- sandhill habitat of Putnam County, north- mately 21 hooks in a half ellipse open to eastern Florida. the rear. A8 with spiracle greatly enlarged, Etymology.-The specific name is de- equalling size of prothoracic spiracle; 10 rived from the Greekpholeter (one who lurks pairs of primary setae, SV bisetose. A9 with in a hole), in reference to its subterranean 9 setal pairs, SV unisetose. Anal plate light behawior. brown, bearing 4 pairs of setae. Discussion. -Acrolophus pholeter does Pupa (Figs. 23-32).-Length of largest not appear closely allied to any North pupa: male, 7.2 mm; female, 11 mm. Light American Acrolophus. In addition to its dis- reddish brown in color. Vertex smooth ex- tinctive male genitalia, this species is un- cept for a pair of minute setae. Antenna and usual in possessing a smooth head and labial palpus of relatively equal length in greatly reduced epiphysis. In color pattern, both sexes; antenna extending to caudal it superficially resembles a nearly unicol- margin of A3 and just short of wings which oms, undescribed species from southern extend to caudal margin of A4; labial palpus Florida and Texas. short. Mesonotum with two pairs of minute Even less can be summarized about larval setae clustered together near midline. Dor- relationships because of the great inadequa- sum of A3-8 with a transverse ridge like cy of our knowledge. Compared with the row of minute spines near anterior margin. few Acroloplzus larvae ever studied (e.g. Da- A9 + 10 relatively smooth except for a clus- vis 1987), the chaetotaxy of A. pholeter ap- ter of 3 pairs of minute spines ventrally and pears little differentiated. However, the a large, slightly bilobed ridge dorsally (Figs. atypical stemmatal reduction in this species, 28-30). particularly the apparent fusion of the three Holotype. -Male, (with associated pupal anterior stemmata, is probably character- exuvium and cocoon) Roberts' Ranch, ca. istic for the species. 6 km north of Hollister, Putnam Co., Flor- Biological observations. -Biology of this ida; em. 12 May 1985, E. G. Milstrey, species has been determined from labora- (USNM). tory and field observations. Apparently, the Paratypes. - FLORIDA: Same data as species is restricted to burrows of the gopher holotype except: 2 8, 3 0, em. Sept. 1984; 5 tortoise, Gopherus polyphemus, in sandhill $,4 n, em. Nov. 1984; 3 a, em. 8 Dec. 1984; habitats (Fig. 1). Tortoise burrows sampled 38, 1 Q,em.19Dec, 1984; 18,m. 5Feb. outside the sandhill biome were found not 1985; 3 9, em. 5 May 1985; 1 a, 9 May 1985; to contain A. pholeter. The sandhills are rel- VOLUME 90, NUMBER 2 177

ict dunes from the Pleistocene and earlier remaining on the body and wings. The poor epochs (Cooke 1945, Laessle 1958). The condition of the adults was not due to the vegetation of this habitat is characterized sampling method but probably due to abra- by longleaf pines, Pinus palustris, several sion occurring during normal adult activity oaks, Quercus laevis and margaretta, with within the sandy burrows. an understory of wiregrass, Aristida stricta, As larval size increased in the burrows, and various herbs. The study site was in density decreased. Larvae were found to be Putnam County, 6 km north of Hollister cannibalistic in laboratory studies. Silk lined (29O41'40"N-8 l048'1O"W). larval galleries were usually constructed just Collections were made every six weeks by below the soil surface in the floor of the vacuum extraction of the burrow using a burrow. Larval galleries could exceed 30 cm modified Echo leaf blower similar to that in length and were often branched. Larvae described by Butler et al. (1984). Larvae, traveled forward and backward in these gal- pupae and adults were obtained but eggs leries, always facing the same direction. were not detected. Larval densities ranged Feeding occurred at the entrance and fecal up to 30 per burrow. In repeatedly sampled pellets were deposited on the soil surface at burrows, population estimates, based upon the other end. removal sampling estimates (Carle and Larvae were successfully reared on go- Maughan 1980), indicated larval numbers pher tortoise fecal pellets. The fecal pellets commonly were 3 to 16. A few burrows not used were mostly partially disgested wire- sampled routinely gave higher estimates, up grass and some oak leaves. Although fecal to as high as 200. First instar larvae were material was available in the burrows, its found from May to November. All other abundance was always low and competition stages were present year round. In the later for it was high. The amount of available part of the summer and early fall the pu- fecal material was generally too low to sup- pating larvae probably were of two gener- port the Acrolophus population present. ations; the previous year's and offspring of Also, larvae were very common in burrows the spring emergence. Larval development that no longer had resident tortoises. In was estimated to require anywhere from 7 active burrows, tortoises frequently re-ex- to 16 months and most likely around 11 cavated their burrows, and in those burrows months (+ one month) for most individu- larval populations were significantly de- als. One lab reared larva, field collected in creased if not exterminated. Larval density its last larval stage, took 15 months to pu- was positively correlated with burrows that pate. The pupation period is relatively short, accumulated leaves and other debris. Ap- normally requiring one to two weeks for parently this rapidly decomposing organic adult eclosion. leaf litter was their primary food source. Pupae and adults were infrequently col- Laboratory studies on limited numbers of lected in the burrows from May to Septem- larvae found that they would survive on the ber. Pupal cocoons (Fig. 5) were constructed litter but that growth rates were lower. of loosely woven silk to which are attached The unique humidity situation in the bur- sand grains and larval frass. Apparently, pu- row appears to be responsible for restricting pation was triggered by unknown narrow the species to this habitat. In the sandhill microclimatic conditions present in indi- gopher tortoise burrows studied, relative vidual burrows, because when pupae were humidities were in the mid to high 90's but found there were commonly more than one, the percent water in the soil was low (0-5O/o) and in other adjacent burrows none were year round. In the laboratory inside an en- found. Adults collected from burrows had vironmental chamber, larvae were found to wings that were badly tattered with few scales survive only within these limits. Mature lar- 178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Carle, F. L. and 0. E. Maughan. 1980. Accurate and vae left their silk wgalleries and moved to drier conditions to pupate. Pupae in the lab- efficient estimation of benthic populations: A comparison between removal estimation and con- oratory could be reared at room tempera- ventional sampling techniques. Hydrobiologia 7 1: ture and normal humidity levels. 181-187. The larvae of the antlion, Glenurus gratis Cooke, C. W. 1945. Geology of Florida. Florida Geo- (Say), and larvae of an undescribed therivid logical Survey Bulletin, 29: 1-339. fly, Arenagena sp. were found to prey upon Davis, D. R., D. H. Clayton, D. H. Janzen, and A. P. Brooke. 1986. Neotropical Tineidae, 11: Biolog- the Acrolophus larvae. Lepidoptera larvae ical notes and descriptions of two new phor- were the only prey either would take under etic on spiny pocket mice in Costa Rica (Lepi- laboratory conditions. Glenurus gratis was doptera: Tineoidea). Proceedings of the probably the major predator. Its larval pop- Entomological Society of Washington 88(1): 98- ulation numbers were more strongly cor- 109. Davis, D. R. 1987. Tineidae, pp. 362-365. In F. W. related with Acrolophus larval numbers. Stehr, ed., Immature . Kendall/Hunt Pub- Both prefer the same burrows and both are lishing Co., Dubuque, Iowa. restricted to the drier, looser sand near the Hubbard, H. G. 1894. The guests of the Florida entrances where the leaves collect; the ther- land tortoise. Insect Life 6(4): 302-3 15. ivid was more ubiquitous. . 1896. Additional notes on the insect guests of the Florida land tortoise. Proceedings of the Entomological Society of Washington 3(5): 299- 302. The senior author (Davis) is responsible . 190 1. Letters from the southwest: Insect fau- for the systematic and morphological por- na in the burrows of desert rodents. Proceedings tions of this paper and the biological ob- of the Entomological Society of Washington 4: 361-364. servations are by the junior author (Mil- Hubbell, T. H. and C. C. Goff. 1939. Florida pocket- strey). Thanks are given to Irwin Roberts gopher burrows and their arthropod inhabitants. for allowing his land to be used for this re- Proceedings of the Florida Academy of Science, search, Judy Gilmore for adult preparation 4: 127-166. Laessle, A. M. 1958. The origin and successional and Dr. Dale Habeck for recognizing the relationship of sand-hill vegetation and sand pine uniqueness of the species and encouraging scrub. Ecological Monographs 28: 361-387. its investigation. We are indebted to Young Milstrey, E. G. 1986. Ticks and invertebrate com- Sohn of the Department of Entomology, mensals in gopher tortoise burrows: Implications Smithsonian Institution, for the line draw- and importance, Pp. 4-1 5. In D. R. Jackson and R. J. Bryant, eds., The Gopher Tortoise and its ings and to Susann Braden and Brian Kahn Community. Proceedings 5th Annual Meeting of of the Smithsonian SEM Lab and Victor the Gopher Tortoise Council. Florida State Mu- Kranz of the Smithsonian Photographic seum, University of Florida, Gainesville, Florida. Laboratory for photographic assistance. The Smith, J. B. 1899. Description of the gopher moth final draft was prepared by Silver West. Canadian Entomologist 3 1: 94-95. Woodruff. R. E. 1982. Arthropods of gopher burrows, pp. 25-49. In R. Franz and R. J. Bryant, eds., The Gopher Tortoise and its Sandhill Habitat. Pro- Butler, J. F., K. H. Holscher, 0. Adeyeye, and E. P. J. ceedings of the 3rd Annual Meeting of the Gopher Gibbs. 1984. Sampling techniques for burrow Tortoise Council. Tall Timbers Research Station, dwelling ticks in reference to potential African Tallahassee, Florida. swine fever virus vectors, p. 1065-1074. In D. A. Young, F. N. and C. C. Goff. 1939. An annotated list Griffiths and C. E. Bowman, eds., Acarology IV of the arthropods found in the burrows of the Flor- Volume 2. Ellis Honvood Ltd. Chichester, En- ida gopher tortoise, Gopherus polyphernus (Dau- gland. din). The Florida Entomologist 22(4): 53-62. Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 2025

Reference Citation: Davis, D. R.; Milstrey, E. G. 1988 [1988.??.??]. Description and biology of Acrolophus pholeter, (Lepidoptera: Tineidae), a new moth commensal from Gopher Tortise burrows in Florida. Proceedings of the Entomological Society of Washington 90:164-178.

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Notes:

File: File produced for the Bibliography of the Neuropterida (BotN) component of the Global Lacewing Digital Library (GLDL) Project, 2005.