taxonID	type	description	language	source
F84F2431FFC5A83BAAFAFA66FBAE9DE4.taxon	type_taxon	Type species Archaeonycteris trigonodon Revilliod, 1917. Grube Messel, near Darmstadt, Germany, Lutetian, early Middle Eocene.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	materials_examined	Holotype HZM 5.38291 isolated left M 3. CL 1.54; TRI W 1.12; TAL W 0.83 (Figs. 1 D – G, 6 D – F). Paratype material HZM 1.31799 right M 1 / 2 in maxillary fragment, metastyle broken CL – CW (protocone to parastyle) 2.34. Protocone lobe (a-p) 1.22 (Figs. 2 A – D, 6 J). HZM 7.38324 left M 1 / 2. CL – CW (protocone to parastyle) 2.50 (Figs. 2 E – H, 6 K – L). HZM 4.37495 right P 2. CL 0.70; CW 0.45 mm (Figs. 1 A – C, 6 G – I). BMNH M 35710 / 12 Trigonid and talonid left M 1 / 2. CL 1.98 (e); TRI W 1.41; TAL W 1.31 (Figs. 1 H – J, 6 A – C). e = estimated measurement.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	distribution	Doubtfully referred material from the type locality HZM 16.31548 part protocone lobe right M 3. (Fig. 2 I – J). HZM 3.35865 protocone lobe right M 1 / 2 (Fig. 2 K – L). Type locality Creechbarrow, Dorset, Nat. Grid Ref. SY 8240 9215, southern England. Stratigraphy Creechbarrow Limestone Formation Bed 12 (Hooker, 1986), Bartonian, late Middle Eocene, Robiacian ELMA. Referred specimen BMNH M 29090, right M 3 talonid; bed F?, Barton Clay Formation, Bartonian, late Middle Eocene; Barton, Hampshire (Hooker, 1986: 241 – 243, fig. 16 f – i).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	etymology	Etymology Latin relicta: alludes to a survivor of an archaic lineage.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	diagnosis	Diagnosis A small Archaeonycteris, mesial CW of M 1 / 2 2.34 – 2.50, CL of M 3 1.54. Protofossa of M 1 / 2 bucco-lingually short, the transverse width subequal with its mesio-distal dimension, instead of clearly exceeding it, as in the three previously described European species listed below. There is no post- or premetaconule crista. Lower molars M 1 – M 3 with the paraconid low, not prominent, and the talonid shorter than the trigonid and with the entoconid virtually absent, forming only a crestiform bulge lingually, lower than the hypoconulid.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	description	Description HZM 1.31799 is a typical Archaeonycteris right M 1 / 2 with no mesostyle, hypocone or conules. The crown is shorter transversely than in A. trigonodon, A. pollex and A. brailloni. The protofossa measured from the protocone cusp to the median point of the centrocrista is shorter than the mesio-distal dimension, instead of clearly exceeding it as in the other described species above. There is a wide and shallow, buccally convex U-shaped centrocrista. The postprotocrista is inserted at the base of the metacone, as in A. pollex, not connected to the metacingulum as in A. brailloni. The crown width of this M 1 / 2 is notably less than in A. brailloni (3.0 – 3.1 mm — Russell et al., 1973: 34). The specimen has the metastyle broken away, but a wide shallow ectoflexus appears to have existed. The resulting ectoflexus is not as deep as in the M 1 of A. brailloni and which is more compatible with M 2 (Russell et al., 1973: fig. 6). The tooth is in situ in a maxillary fragment, which has a small nutrient foramen penetrating the palate below the preprotocrista. F H E G The primary cusps, protocone, paracone and metacone, are conical. The tip of the metacone is damaged mesio-buccally. No small crests emanate from the base of the paracone or metacone as in some specimens of A. brailloni (Russell et al., 1973: 30) and there is no trace of any pre- or post-paraconule crista or of the paraconule or metaconule on the pre- and postprotocrista. The protofossa is short and deep. The preprotocrista passes mesial to the paracone towards the parastyle. There is no hypocone, but a distinct angularity at the disto-lingual base of the protocone. The parastyle is massive, with a bluntly rounded tip projecting mesially. The apparent absence of any buccal cingulum or stylar cusps, which are well developed in A. brailloni could be due to enamel erosion evident on the buccal margin of the tooth. The metastyle is missing, but it is clear that the buccal edge was recessed, without any trace of a mesostyle. The protocone is surrounded lingually by a wide cingulum, which terminates abruptly on the mesial border of the crown, just in front of the level of the protocone tip. Distally the cingulum forms a straight border to the protocone lobe and is angulated outwards at the base of the metacone, where it is interrupted by a crack in the specimen. There is no evident internal distortion of the metacone base resulting from the crack. The tooth was originally threerooted, of which only the robust mesio-buccal root remains intact. HZM 7.38324 (Figs. 2 E – H, 6 K – L) is a left M 1 / 2, lacking the metacone and metastyle, which are broken off. The protocone and paracone are low, tubercular cusps, virtually unworn. The strong preprotocrista extends via the paracingulum to the blunt, broad parastyle, with a rounded mesiobuccal angularity. The paracone is slightly lower than the protocone and the preparacrista ends buccally with a small stylocone. The protofossa is deep and like HZM 1.31799 it is relatively short transversely. Judging from the area of its broken base the missing metacone was smaller than the paracone and there is no hypocone. The protocone is slightly slanted mesially. The preserved part of the ectoflexus is mesially positioned. It is shallower than in A. brailloni. A robust cingulum surrounds the protocone base. Distally it is broken off close to the metacone; mesially it extends buccally to join the paracingulum. The buccal surface of the paracone is concave. A broad buccal cingulum is present, broken off at the same level as the centrocrista. Three roots were originally present. HZM 16.31548 (Fig. 2 I – J) comprises the mesial half of the protocone of a right M 3 with a strong mesial cingulum, inserted below the preprotocrista in a similar position to HZM 1.31799. The protocone is slightly more curved buccally than in the latter specimen. The paracone has been obliterated. The specimen is considered to be part of an M 3. HZM 3.35865 (Fig. 2 K – L) is a fragment of the protocone of a right M 1 / 2. Apart from the strong cingulum only the high protocone cusp and part of the preprotocrista are preserved. It appears distinctly larger than HZM 1.31799, but has a very similar shallow hollow between the preprotocrista and the robust cingulum. HZM 4.37495 is a right P 2 (Figs. 1 A – C, 6 G – I), resembling in most respects this element in A. brailloni (Russell et al., 1973). Although the mesially situated cusp tip is worn, it is clearly low-crowned, with its buccal border quite strongly convex and the lingual border almost straight. A mesial cristid curves lingually to the crown tip. A distal cristid is almost straight and is evidently situated more lingually than in A. brailloni. In this specimen the talonid is slightly narrower than in A. brailloni and distinctly upturned. A relatively very strong cingulum surrounds the crown, eroded distobuccally. The single robust root is angulated distally below the crown. It may be noted here that the lower premolars of Stehlinia are high crowned and sharply point- ed. Small single-rooted anterior lower premolars resembling this in shape are not present in hipposiderids or rhinolophids. M 35710 / 12 is a left M 1 / 2 talonid and trigonid described and figured by Hooker (1986: 242, fig. 16 a – e). Found in the same hole, they were judged to belong to the same specimen, although the fit is not perfect. The specimen, remeasured from a cast after repositioning of the two fragments, gives a revised CL 1.98; TRI W 1.41; TAL W 1.31 (Figs. 1 H – J, 6 A – C). HZM 5.38291 holotype (Figs. 1 D – G, 6 D – F) is an intact left M 3. It resembles in most respects the M 1 / 2 M 35710 / 12, but the more reduced talonid shows that it is an M 3. It is well preserved, apart from the broken paraconid and slight enamel loss at the distolingual base of the crown. Only the cusp tips of the protoconid and metaconid are lightly worn. It is similarly a low crowned tooth, with the talonid distinctly shorter than the trigonid. The protoconid is dominant, the metaconid rather lower and the eroded paraconid appears to have been lower and less prominent than in the M 1 / 2 above. In the talonid the hypoconid is distinct, but low. The hypoconulid is almost central, projecting distally slightly less than the M 1 / 2. The entoconid is virtually absent, forming only a slight crescentic bulge disto-lingually. The talonid is distinctly more reduced than that of A. brailloni M 1 / 2 (Fig. 3 C – D). A strong continuous buccal cingulum extends from the precingulid to the postcingulid. It is less angulat- ed below the protoconid than that of the M 1 / 2. No lingual cingulum is present. The short cristid obliqua is inserted almost centrally below the trigonid notch and it is notched near its junction with the trigonid. The mesial root of the tooth is preserved, the distal one has broken off. Hooker (1986: 242, fig. 16 f – i) described a right M 3 talonid (M 29090) TAL W 0.93 from the Barton Clay (? bed F) sharing a number of features with the Creechbarrow M 1 / 2 and the M 3 described here. Comparison of this Barton Clay talonid with the holotype M 3 above shows that it is identical in structure and it is here confidently referred to A. relicta. Comparisons Most of the teeth of Archaeonycteris recovered are fragmentary. HZM 1.31799, the most complete M 1 / 2 so far found differs from previously described species in having a much smaller protofossa. This condition is linked with smaller size of the talonid of the lower molars and with the reduced entoconid. This nearly intact M 1 / 2 exhibits three of the diagnostic characters of Archaeonycteris (Russell and Sigé, 1970: 125). 1. Absence of the mesostyle. This is accompanied by a deep ectoflexus, listed by Simmons and Geisler (1998: 42) as the first of the apomorphies distinguishing Archaeonycteris. 2. Absence of the hypocone. 3. Absence of the metaconule, although a small paraconule may be present (Russell and Sigé, 1970: 125). The conules are small or absent. HZM 1.31799 (probably an M 2) and HZM 7.38324 are less transversely elongated than in all three European species, with a distinctly smaller protofossa and shallow and U-shaped centrocrista. The apparent absence of buccal cingulum and cingular cusps on the parastyle of HZM 1.37199 may be, at least in part, due to erosion of the enamel. This tooth resembles A. brailloni (Fig. 3 A – B) in having an indication of a hypoconal lobe, absent in HZM 7.38324; this feature is not constant in A. brailloni (Russell et al., 1973: fig. 6). It resembles A. pollex in having the post-protocrista directly inserted on the base of the metacone, rather than continuing into the metacingulum as in A. trigonodon and A. brailloni. The lower molars of A. relicta have reduced talonids, with the entoconid virtually absent. Comparison is necessary with M 1 / 2 of some other genera of European Early and Middle Eocene Chiroptera and recently described Indian archaeonycterids. Ageina tobieni Russell, Louis and Savage, 1973 is a taxon of uncertain familial status, most recently referred to Nataloidea incertae sedis (Simmons and Geisler, 1998: 134, table 8). The M 1 / 2 show some similarity to the Creechbarrow M 1 / 2 (HZM 1.31799) in having a rather subquadrate basal outline, an ectoflexus, absent mesostyle and prominent para and metastyles (Russell et al., 1973: fig. 8). It is clear, however, that this form differs in having a sharply angulated centrocrista. The trigon basin of the Creechbarrow M 1 / 2 is distinctly smaller and rounded buccally. The lingual cingulum is incomplete at the base of the protocone in Ageina tobieni, strong and complete in the Creechbarrow A. relicta. (see below for comparison of Ageina with an unidentified Microchiropteran trigonid). Stehlinia Revilliod, 1919. Bats of this genus, referable to the Nataloidea according to Simmons and Geisler (1998: 136, table 8) have an M 1 / 2 with a pronounced mesostyle and small crests originating from the bases of the paracone and metacone. Icaronycteris? menui Russell, Louis and Savage, 1973 has the M 1 / 2 more transversely elongate in form and also has the centrocrista sharply angulated. The lingual cingulum is variably complete (Russell et al., 1973: fig. 4) Palaeochiropteryx Revilliod, 1917. Bats of this genus have a well developed mesostyle and the ectoflexus is shifted mesially with virtually absent concavity between the mesostyle and metastyle, as well as a sharply angulated centrocrista (Russell and Sigé, 1970: figs. 4, 6 and 7, plate 1). Protonycteris gunnelli Smith, Rana, Missaien, Rose, Sahni, Singh, and Singh, 2007 was recently described from the Ypresian of Vastan Lignite Mine, Gujarat, India (Smith et al., 2007: 1004), based on the unique holotype left dentary, with P 3 – M 3 in situ. It is slightly larger than A. relicta (M 1 CL 2.00; TRI W 1.40; TAL W 1.40; M 2 CL 2.10; TRI W 1.50; TAL W 1.40; M 3 CL 1.90; TRI W 1.35; TAL W 1.05). It resembles A. relicta in having a reduced entoconid, but differs in having a distobuccally incomplete ectocingulum and short weak cristid obliqua in the lower molars. The talonid of M 3 is longer and the hypoconulid is shifted lingually. The maxillary dentition is as yet undescribed. Archaeonycteris? storchi described from the same Indian Early Eocene site (Smith et al., 2007: 1007) and known from a left dentary fragment with M 1 – M 3 in situ is rather smaller in size than A. relicta (M 1 CL 1.50; TRI W 0.95; TAL W 1.00; M 2 CL 1.55; TRI W 1.05; TAL W 1.00). It has a more distinct and relatively stronger entoconid, the less reduced talonid thus differing strikingly from A. relicta. The maxillary dentition remains undescribed and its inclusion in Archaeonycteris remains uncertain. Archaeonycteris? praecursor Tabuce, Antunes and Sigé, 2009. This taxon has been recently described from a single right M 1 / 2 (UNLSNC — 447) from the Early Eocene of Silveirinha, Portugal. This tooth, with a CL 1.45 and CW 0.95 is slightly small- er than the two lower molars at present known of A. relicta and differs strikingly in morphology. Its cusps are contrastingly tall and pointed, with a more prominent, mesially directed paraconid. The entoconid is furthermore well developed, subequal in height with the hypoconid; the hypoconulid is more prominent and high, sublingual in position (Tabuce et al., 2009). Whatever the ultimate generic status of this bat may prove to be, it is clearly quite different from A. relicta. Rather few measurements are available for the lower molars of Archaeonycteris. CL of M 1 – M 2 2.40 – 2.48 (n = 4, m = 2.45); M 3 CL 2.12 – 2.20 (n = 2, m = 2.16) in A. pollex; CL M 1 – M 2 2.00 – 2.20 (n = 4, m = 2.09); M 3 1.88 (n = 1) in A. trigonodon (Storch and Habersetzer, 1988: table 2); A. brailloni M 1 CL 2.30 (n = 1). These measurements (crown widths not available) suggest that the lower molars of A. relicta are smaller, consistent with the reduced talonids observed. In A. pollex the relatively large hypoconulid is very high, as high as the entoconid or higher (Habersetzer and Storch, 1988). In A. brailloni the talonid cusps are all better developed than in A. relicta with the entoconid only somewhat reduced (Fig. 3 C – D).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFC5A83EA9B9FE68FC019BA7.taxon	discussion	Discussion This is the first confirmed occurrence of Archaeonycteris in the British fossil record. Savage and Russell (1983) recorded ‘ Archaeonycteris ’ from the Bartonian of Robiac but all other records of the genus are from older rocks. Legendre et al. (1992: 421, fig. 7 a – b) figured an archaeonycterid of undetermined genus and species from Vielase, Phosphorites du Quercy, Early Eocene. Their figured upper molar (fig. 7 a) has a transversely elongated trigon basin quite unlike A. relicta. The occlusal view of the figured M 1 / 2 shows an entoconid better developed than in A. relicta. Discovery of more intact material from the British Bartonian is needed to allow comparison of other tooth types.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCFA833AAA5FD69FDA59DE5.taxon	materials_examined	Lectotype NMB QP 875 right maxilla with C 1 – M 3, selected by Sigé (1978: 253) from the Phosphorites du Quercy. Material HZM 1.31792 left M 2. CL 1.79; CW 2.11 (Figs. 4 A – B, 7 A – B). HZM 3.38170 left M 1 / 2. CL 1.82; TRI L 0.96; TRI W 1.25; TAL L 0.83; TAL W 1.28 (Figs. 4 C – F, 7 C – E). Doubtfully BMNH M 36790 basal part left C 1. CL 1.77; CW 1.53 (Fig. 9 A – C).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCFA833AAA5FD69FDA59DE5.taxon	description	Description The left M 2 (HZM 1.31792) conforms in size with the assemblage from Ste-Néboule described by Sigé (1978: table 2) in which 40 examples of isolat- ed M 2 measured 1.6 – 1.87 mm in crown length and 1.94 – 2.21 mm in crown width. The morphology of the Creechbarrow specimen closely resembles the M 2 in a maxilla (BRE 1 - 775) from Le Bretou (Sigé, 1988: 91, fig. 27). Although the postcingulum of the Creechbarrow tooth has been eroded the shape of the protocone lobe is more comformable with M 2 than M 1. Other important morphological points are the mesially curved parastyle and shorter straight metastyle; the ectoflexus is very shallow; the postprotocrista stops at the lingual base of the metacone as a result of which the protofossa is enclosed, narrow and deep as described by Sigé (1988: 90). This tooth has been compared with a cast of the lectotype (Sigé, 1978). The M 2 from Creechbarrow has identical dimensions to the lectotype M 2 and differs only in having a less pointed mesostyle. Pseudorhinolophus morloti (Pictet, 1857) is smaller (M 1 CL 1.42 – 1.56; M 2 CL 1.32 – 1.55, M 1 / 2 CL 1.39 – 1.62 (Hooker and Weidmann 2000: table 11); M 1 / 2 have prominent lingually projecting hypocones (Hooker and Weidmann, 2000: fig. 37 d). Pseudorhinolophus weithoferi Revilliod, 1917, is larger. The M 2 in a cast of the syntype (NMB QH 192) measures CL 2.43; CW 2.56. M 36790 is the basal part of an upper left canine, described and figured by Hooker (1986: 245, fig. 17 e – g). Its dimensions are very slightly larger than the series of eight C 1 from Le Bretou listed by Sigé (1988: table 4): CL 1.55 – 1.72; CW 1.18 – 1.36. It is very tentatively placed here because its morphology resembles that of Pseudorhinolophus. HZM 3.38170 (Figs. 4 C – F, 7 C – E) is a well preserved left M 1 / 2 with moderate wear of the cusp tips and crests, especially the paracristid, postmetacristid and metaconid, with exposure of dentine. The crown outline is subrectangular with its width greater in relation to its length than in HZM 1.31222 described below as Rhinolophidae gen. et sp. indet. The paracristid is slightly convex mesially, the metacristid almost straight. Both crests lack the pronounced angulation present in HZM 1.31222, possibly due to the more advanced wear in this specimen. The blunt paraconid projects much more lingually than in HZM 1.31222, producing the distinctly rectangular outline. The pattern of wear on the lingual aspect of the trigonid and talonid resembles that seen in the modern Hipposideros larvatus (HZM 23.35012: Kyway Chai Village, Rakhine State, Myanmar) and it is noteworthy that the small cingular ledge beneath the lingual opening of the trigonid basin seen in HZM 1.31222 is absent in this specimen as well as in the modern hipposiderid. The cristid obliqua, also worn with exposure of dentine, is attached low down on the distal trigonid wall and more distinctly lingual to the midline than in HZM 1.31222. The hypoconulid is situated buccally of the entoconid, both cusps exhibiting slight wear. The pre-entocristid is also worn, forming a narrow marginal gutter on the lingual border of the talonid, which is distinctly wider than the trigonid. The talonid is low with the hypoconid about two-thirds of the protoconid height. A well preserved narrow cingulum extends from its origin close to the paraconid around the buccal and distal crown margins to the hypoconulid. It is not robust as in Archaeonycteris and in Stehlinia and lacks the marginal crenulations present in the latter. There were two transverse roots, the distal one preserved and direct- ed slightly distally. Good occlusion can be demonstrated with HZM 1.31792, considered to be an M 2 of this species. The F measurements of this specimen fall within the range of the series of 10 M 1 / 2 from Le Bretou listed by Sigé (1988: table 4). CL 1.62 – 1.82 (m = 1.70); CW 1.2 – 1.49 (m = 1.33, n = 10) and of 60 M 1 / 2 from Ste- Néboule; CL 1.76 – 1.95 (m = 1.84); CW 1.25 – 1.59 (m = 1.43) (Sigé, 1978: table 2).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCDA832AB07FE6AFE819BA7.taxon	materials_examined	Material HZM 1.31222 left M 1 / 2: CL 1.95; TRI W 1.09; TRI L 1.15; TAL W 1.15 (e); TAL L 0.77 (Figs. 4 G – J, 7 F – H). HZM 2.38159 trigonid fragment of right lower molar TRI W 1.47; TRI L 1.15; PH 1.63 (Fig. 8 D – F). BMNH M 35711 trigonid fragment left lower molar TRI W 1.17 (Fig. 8 A – C).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCDA832AB07FE6AFE819BA7.taxon	description	Description The left M 1 / 2 (HZM 1.31222) clearly belongs to a different taxon from the specimens here referred to Pseudorhinolophus schlosseri. It has a more elongated and narrower outline, with the cusps and crests more gracile. The paraconid is directed more mesially, thus lacking the subrectangular outline of P. schlosseri M 1 / 2. It is a well preserved tooth, intact except for slight erosion of the basal crown edge especially below the entoconid. The trigonid is longer than the talonid, measured by the method of Sigé (1968). The cusp tips are lightly worn; the cusps of the trigonid are prominent, the protoconid dominant, the metaconid just higher than the upturned paraconid. The cristids are sharply angulated at the protoconid. Beneath the more widely open trigonid basin there is a distinct short lingual cingulum, which is absent in the specimen of P. schlosseri. The distal wall of the trigonid is vertical and high. The talonid is judged wider than the trigonid despite basal erosion. The hypoconulid is prominent, its cusp directed upwards and it is situated slightly buccally to the entoconid, but closer to this cusp than the hypoconid. The preentocristid declines steeply to form a rounded talonid notch extending below the level of the cristid obliqua, contrasting strongly with Stehlinia in this respect. It is lingually concave, thus conforming with Vaylatsia prisca (Revilliod, 1920). The unworn cristid obliqua is attached more buccally to the posterior trigonid wall than in P. schlosseri and close to the midline. The buccal cingulum was continuous from its origin just distal to the base of the paraconid to the base of the hypoconulid, but is partly eroded. It is attached closer to the tip of the paraconid than in Stehlinia. It is compatible in size with the M 2 of P. schlosseri described above, but the two teeth do not occlude well with each other, supporting the conclusion that this M 1 / 2 belongs to a different taxon. HZM 2.38159 and M 35711 are isolated trigonids similar in size and morphology to HZM 1.31222 described above. Their mesial cingulum extends across the trigonid notch. These trigonids are slightly less open than in the intact specimen, perhaps indicating that they are M 2 ’ s and the intact specimen is an M 1.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCDA832AB07FE6AFE819BA7.taxon	discussion	Discussion In the absence of complete mandibular and maxillary dentitions considerable difficulty is bound to be encountered in identification of isolated rhinolophoid teeth. The marked morphological similarity of the two specimens here referred to P. schlosseri to the lectotype and other published material has enabled a confident identification of this species. However, this second M 1 / 2 (HZM 1.31222) is a much more complicated and difficult problem. The genus Palaeophyllophora Revilliod, 1917 can be excluded because the talonid of M 1 – M 2 is much reduced in this genus relative to the trigonid (Sigé, 1978: 255, plate 1, fig. 4). The hypoconulid is stronger and less closely situated to the entoconid, more nearly median in position. Genus Pseudorhinolophus Schlosser, 1887 has been consider- ed a subgenus of Hipposideros Gray, 1821 by some authors (e. g., Sigé, 1978: 251; 1988: 89). The taxonomic history of P. schlosseri has been reviewed by Sigé (1978). This species is represented by abundant material from the Late Eocene of Ste-Néboule, including 60 specimens of M 1 – M 2 and from the late Middle Eocene of Le Bretou (10 M 1 – M 2) listed by Sigé (1988). Morphological differences from HZM 3.38170 have been noted above, which suggest that HZM 1.31222 from Creechbarrow may belong to a different genus from Pseudorhinolophus. Le Bretou has also yielded an unidentified rhinolophoid taxon, Rhinolophoidea sp. indet. of Sigé (1988: 93). This form also has morphological features suggestive of Rhinolophus. It is clear however, that the Creechbarrow tooth cannot belong to the same taxon, as that from Le Bretou, since the Le Bretou material of M 1 – M 2 is smaller CL 1.39 – 1.67 (m = 1.57); CW 1.02 – 1.39 (m = 1.20). It is also clear from Sigé’s (1988: 94, fig. 32) description that the M 1 / 2 of Le Bretou is less narrow in relation to its length and does not have the pre-entocristid so steeply declining. The possibility therefore exists that the Creechbarrow Rhinolophidae indet. could be referable to genus Rhinolophus Lacépède, 1799 or Vaylatsia Sigé, 1995. The latter genus is represented in the Eocene by V. prisca (Revilliod, 1920), which is similar in size to P. schlosseri (Sigé, 1978: 261) and morphologically rather similar. Unfortunately the exact provenance of much of the material of the two species attributed to Rhinolophus by Revilliod (1920), and hence their age is unknown. Hooker and Weidmann (2000: fig. 38 a – c) have selected and illustrated the lectotype of Rhinolophus priscus, a left dentary with P 4 – M 3, slightly distorted and alveoli of C 1, P 2 – P 3 (NMB Mt 992) from Entreroches. It is the only specimen from Revilliod’s (1920) syntypes with a precise locality. Comparison of HZM 1.31222 with a cast of the lectotype reveals considerable similarities. However, the molars of the lectotype are not in good condition making identification problematic. Rhinolophus pumilio Revilliod, 1920 is a smaller species, similar in size to Pseudorhinolophus morloti, of uncertain stratigraphic origin in Le Quercy. Sigé (1995) considered the genus Vaylatsia to belong to the Hipposideridae. Dentally the genus resembles Rhinolophus but with distal humeri like hipposiderids. Hooker and Weidmann (2000: 60) have, however, pointed out that the hipposiderid humeral structure is primitive for the Rhinolophoidea and that Vaylatsia may therefore be better placed in the Rhinolophidae. HZM 1.31222 certainly exhibits some distinct similarities to Recent Rhinolophus as exemplified by R. rouxii (HZM 72.36548 from Tirunelveli, Tamil Nadu, India). This species also has gracile cusps and crests, a rather mesially directed paraconid, a small cingular ledge below the trigonid basin opening and a steeply sloping preentocristid. Comparison of a variety of Recent species of Rhinolophus and Hipposideros suggests that the dental differences of HZM 1.31222 listed above are distributed in both genera in a random manner partly related to size. It is, however, clear that a relatively narrow M 1 / 2 in relation to its length, especially with gracile crests and cusps as in this specimen is much more likely to be rhinolophid than a hipposiderid. Unequivocal identification as rhinolophid would require the finding of a jaw with the alveolus of a small vestigial, labially situated P 3 (Sigé, 1988: 93). This vestigial tooth is, however, not always present in Rhinolophus.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCCA835A940FDBFFBF69DDA.taxon	materials_examined	Holotype Right mandibular ramus with C 1 – M 3, ascending ramus broken. Phosphorites du Quercy, NMB QP. 727. Material HZM 4.36362 right C 1. CL 1.34; CW 0.96; CH (labial) 1.86 (base of cingulum to tip); total length with root 3.33 (Figs. 5 A – D, 9 D – F).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCCA835A940FDBFFBF69DDA.taxon	description	Description HZM 4.36362 is a well preserved lower right canine of a medium small microchiropteran, with the root intact. The crown is relatively short and broad at the base, with the tip curved lingually. The distal surface has a central ridge bordered on each side by shallow grooves; on the lingual surface there is a mesial groove, while the buccal surface is quite smooth and evenly convex. The lower canine of S. quercyi described here agrees well with the description of this element in Stehlinia minor Revilliod, 1922 provided by Sigé (1974: 260, fig. 2) differing only in being significantly larger and in having a pronounced median ridge, bordered by grooves on its distal surface. A distinctive feature of HZM 4.32362 is the presence of a mesio-lingual cingular cusp, which is lacking in the lower canine of Archaeonycteris (Fig. 3) as well as in the C 1 of modern Rhinolophus and Hipposideros species. It is also absent in the C 1 of Vespertiliavus gracilis Revilliod, 1920 (Sigé, 1988: fig. 8) The C 1 of P. schlosseri is distinctly shorter (CL 1.06 – 1.18 but wider CW 1.05 – 1.13). Sigé (1988: table 4) showing that this is a much more gracile tooth. It also lacks the mesio-lingual cingular cusp.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCCA835A940FDBFFBF69DDA.taxon	discussion	Discussion Three species of Stehlinia occur in the European Tertiary. All three are now known to occur in Britain. Stehlinia minor (Revilliod, 1922) is record- ed from the Late Eocene Bembridge Limestone Formation, Headon Hill (Hooker et al., 2004: 163). Stehlinia quercyi is a distinctly more robust species recorded here for the first time in Britain from the Creechbarrow Limestone. Stehlinia gracilis Revilliod, 1919 occurs in the Early Oligocene Hamstead Member, Bouldnor Cliff, Isle of Wight (Hooker et al., 2004) Although only represented by a canine, size, the main criterion for distinguishing species of Stehlinia, determines the presence of S. quercyi at Creechbarrow. This species, apart from the essentially unlocalised holotype (Quercy undifferentiated), is recorded from Perrière (Quercy) (Remy et al., 1987) and Eclépens B, Switzerland (Hooker and Weidmann, 2000). Both records are from the early Priabonian (early Headonian ELMA). The Creechbarrow record therefore represents a range extension back into the Bartonian.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCBA836A98CFE76FE689DC0.taxon	materials_examined	Material HZM 22.34774 trigonid right lower molar: TRI W 0.77; TRI L 1.66; PH 1.28 (Figs. 5 E – H, 8 G – I).	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCBA836A98CFE76FE689DC0.taxon	description	Description This is a small gracile right trigonid. It is certainly different from all the Creechbarrow taxa described above. The cusps are completely unworn, so that their relative height is clearly determined. The protoconid is dominant, the metaconid high, attaining three-quarters of its height in distal view; the paraconid is lowest, connected to the protoconid by a shallow concave paracristid. Distally there is a distinct median trigonid notch, with the attachment of 2 the cristid obliqua directly below it and at a lower level than the broken off pre-entocristid lingually. The robust buccal cingulum is broken off just distal to the protoconid and extends to a point buccal to the paraconid. There are no crenulations visible on it, but a shallow concavity is present midway along its mesial border. The trigonid basin is widely open and the lingual aspect of the protoconid has a distinct vertical ridge (bordered on both sides by hairline cracks), and with distinct hollows both mesial and distal to the ridge, which passes to the base of the metaconid.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
F84F2431FFCBA836A98CFE76FE689DC0.taxon	discussion	Discussion It does not resemble the trigonids of modern emballonurids compared or of Vespertiliavus. It is equally incompatible with Recent small molossids or with Kerivoula hardwickei, all of which have notably lower paraconids as well as other differences. The closest similarity to a described bat is with Ageina tobieni Russell, Louis and Savage, 1973 from the Early Eocene of Mutigny, France. The right M 1 M. N. H. N – Mu 5112 shares with the Creechbarrow trigonid the presence of a lingual protoconid rib (not shown in their fig. 8 f) passing to the metaconid and it appears compatible in size and general morphology. Intact teeth are needed, since this small species may possibly represent Ageina; meanwhile its identity is obscure. Ageina remains of uncertain familial affinity.	en	Harrison, David L., Hooker, Jeremy J. (2010): Late Middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, southern England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12 (1): 1-18, DOI: 10.3161/150811010X504554
