PUBLICATIONS OF THE MUSEUM TEXAS TECH UNIVERSITY

Three publications of The Museum of Texas Tech Univer¬ sity are issued under the auspices of the Dean of the Gradu¬ ate School and Director of Academic Publications, and in cooperation with the International Center for Arid and Semi- Arid Land Studies. Short research studies are published as Occasional Papers whereas longer contributions appear as Special Publications. Papers of practical application to col¬ lection management and museum operations are issued in the Museology series. All are numbered separately and pub¬ lished on an irregular basis.

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ISSN 0149-175X

Texas Lech Press Lubbock, Texas 79409

OCCASIONAL PAPERS

THE MUSEUM

TEXAS TECH UNIVERSITY

NUMBER 102

26 AUGUST 1986

THE GREATER HORSESHOE BAT, RHINOLOPHUS FERRUMEQUINUM (SCHREBER, 1774), IN GREECE, WITH DESCRIPTION OF A NEW SUBSPECIES

Joan Iliopoulou-Georgudaki and John C. Ondrias

The greater horseshoe bat, Rhinolophus ferrumequinum, is widely distributed in Europe, Asia Minor, Persia, and the Arabian Peninsula, eastward through southern Asia to Japan. In North Africa, it is known from Algeria and Morocco and may occur elsewhere on that continent as well. A number of subspecies have been described (Felten et ai, 1977), of which the most acceptable seem to be R. /. ferrumequinum (Schreber, 1774), R. /. martinoi Petrov, 1941, and R. /. proximus Andersen, 1905. The recognition of other named races is questionable.

The species has been recorded from Greece from the following localities: Parnassos and Syros (Miller, 1912); Rhodos (de Beaux, 1928; Festa, 1914); Olympos (Chaworth-Musters, 1932); Skyros (Pohle, 1953); Kimmeria (Thrace) (Linderg, 1955); Corfu (Niethammer, 1962); Ioannina (Epirus) (Felten et ai, 1977); Crete (Bate, 1906; Felten et al., 1977; Kahmann, 1959; Miller, 1912; Pieper, 1977).

Materials and Methods

The present study is based on examination of 120 specimens from continental Greece and 17 from ihe Island of Crete. The localities of collection are listed in the accounts beyond. Most specimens are preserved in alcohol and are deposited in the Zoological Museum of the University of Patras (ZMUP), Greece.

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OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY

Table 1. — Body and skull measurements of Rhinolophus ferrumequinum from the Island of Crete and North, Central, and South Greece, with i-values comparing Cretan samples with each of Grecian ones. Significant t- values (P*^

0.01) are indicated by an asterisk.

Char -

aciers

Crete

North Greece

Central Greece

South Greece

N

Mean

SD

N

Mean

SD

/-values

N

Mean

SD

(-values

N

Mean

SD

(-values

TL

13

99.30

3.16

8

110.38

4.50

6.37*

57

106.70

4.84

5.20*

44

104.75

3.71

4.72*

Ef B

13

62.00

2.76

8

70.19

2.48

6.85*

55

67.01

2.01

5.63*

44

65.22

2.74

3.72*

T

13

37.30

2.12

8

40.19

4.74

1.86

55

39.23

3.66

1.80

44

39.53

2.73

2.64

HE

13

10.61

0.85

8

1 1.34

0.56

2.84

57

11.16

0.64

2.62

44

1 1.01

0.49

3.02*

E

13

22.16

1.60

8

25.68

0.76

5.31*

61

23.81

1.08

3.74*

44

22.47

3.20

0.01

FA

15

55.51

1.64

8

58. 1 6

0.64

4.32*

61

57.71

1.29

5.60*

42

58.14

1.40

5.99*

(.l.S

10

21.63

0.27

7

22.69

0.30

7.17*

54

22.60

0.39

7.51*

27

22.43

0.38

6.00*

Cl.

10

19.55

0.45

7

20.5 1

0.30

4.91*

57

20.37

0.32

7.01*

27

20.21

0.32

3.86*

ZB

1 1

12.10

0.26

7

12.54

0.25

3.50*

60

12.41

0.26

3.64*

34

12.31

0.24

2.76*

BB

12

8.82

0.29

7

9.06

0.24

1.85

59

9.10

0.22

3.80*

32

9.12

0.24

3.42*

PC

1-1

2.59

0.16

7

2.67

0.19

1.00

62

2.66

0.15

1.56

38

2.62

0.18

0.59

CM3

11

7.97

0.12

7

8.38

0.23

5.93*

63

8.38

0.19

7.72*

42

8.35

0.16

7.69*

MIR

11

8.98

0.14

7

9.25

0.12

5.27*

58

9.45

0.22

7.61*

42

9.11

0.20

7.32*

M

13

15.31

0.26

7

15.98

0.24

5.72*

59

15.89

0.28

6.84*

39

15.81

0.27

6 00*

Key to characters: TL, total length; HB, length of head and body; T, length of tail; HE. length of hind foot; E'. length of ears; E'A, length of forearm; GLS, greatest length of skull; CL, condylobasal length, ZB, zygomatic breadth; BB, breadth of braincase; PC, postorbital constriction; CM3, maxillary toothrow; MIR,

mandibular toothrow; M, length of mandible.

All measurements are recorded in millimeters. Color variation was analyzed by comparing skins of similarly aged animals directly with the color plates of Ridgway (1885). For each specimen, external and cranial measurements, sex, and date and locality of collection were recorded. Only adults, as determined by complete ossification of the phalanges and by at least some wear on the teeth, were included in statistical treatment of mensural data.

Initial analysis involved computation of standard statistics (mean, range, standard deviation, and standard error of the mean) for 14 variables of all specimens from Greek localities. The same metric data were subjected also to a /-test (fable 1). Morphological data from each of the comparable samples were analyzed using a computer program that generated coefficients of average taxonomic distance (ATD) among samples from standardized character values, as well as by the unweighted pair- group method using arithmetic averages (UPGMA) based on the distance coefficient (Fig. 1). A data matrix was formed using localities as OTU’s (Operational Taxonomic Units) and size characters as variables. The distance matrices and dendrograms facilitated interpretation of overall similarities between OTU’s.

ILIOPOULOU-GEORGUDAKI AND ON DR IAS — HORSESI l()l BA I

3

10.

9-

Yu

C.Gr

Cz

C rete

8 -

Fig. I. Phenogram computed from distance matrix based on standardized characters and clustered by unweighted pair-group method using simple averages (UPGMA). Yu, Yugoslavia; C.Gr, central Greece; Cz, Czechoslovakia.

Results

Sexual variation. — YVe detected no statistically significant sexual differences in measurements or coloration in any of the available samples. Consequently, data relating to both sexes were combined in subsequent analyses.

Age variation. — We noted no significant differences among adults in the several samples except increased wear on teeth with age.

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OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY

Seasonal variation. — At localities from which specimens were available from more than one time of the year, a slight seasonal variation in color was discernible. Thus, in specimens from Peloponnesus we found darker shades of brown in the winter (intermediate between hair brown and drab) than were found in the spring (natal brown).

Geographic variation. — There is variation in color and mensural values among samples. Evidence of such variation is depicted in Fig. 2, in which length of forearm is compared between samples from mainland Greece and Crete. Furthermore, the color of individuals in samples from continental Greece is a darker shade of brown from south to north. Specimens from Crete are noticeably paler and more brownish than are those from continental Greece, verona and benzo brown in winter.

Discussion and Taxonomic Assessment of Samples

For taxonomic assessment of Greek populations (especially for the one on Crete) of R. ferrumequinum , we compared samples from four geographic areas, each within the range of a single subspecies as currently recognized: Czechoslovakia (Hanak, 1964), Yugoslavian Macedonia (Miric, 1960), continental Greece, and the Island of Crete. Smaller samples from separate Greek localities have been incorporated into larger categories representing the northern, central, and southern parts of the country.

Czechoslovakia is occupied by R. ferrumequinum ferrumequi¬ num and southern Yugoslavia and continental Greece by R. ferrumequinum martinoi, whereas the Island of Crete is occupied by a new subspecies described in this paper. Average measurements for the four samples are presented in Table 2.

On the basis of nine cranial and external measurements the sample from Crete is separable from the samples of R. f. ferrumequinum and R. f. martinoi by an average taxonomic distance of 8.22. A dendrogram indicating levels of morphologic difference, based on the largest sample, that from central Greece, places the population from Crete as distinct at the subspecific level (Fig. 1). These differences as well as differences between specimens from Crete and continental Greece, as concluded by the /-test, lead us to propose the following taxonomic arrangement of Greek specimens of R. ferrumequinum.

ILIOPOULOU-GEORGUDAKI AND ON DR IAS — HORSES HOF BA 1

62 .

61

Peloponnesus N = 42

60

59

58

57.

56

55

54

Crete

N=15

Sterea Hellas Nz36

N. Greece N = 8

53.

Fig. 2. Geographic and individual variation in length of forearm in four Greek samples of Rhinolophus ferrumequinum. Ihe har represents one standard deviation on either side of the mean (horizontal line) in each sample, whereas the vertical line indicates the extremes in variation.

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OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY

1'able 2. — Means for the specimens of R. f. creticum (Crete), R. I. martinoi ( Central Greece, Yugoslavia ) and of R. f. ferrumequinum ( Czechoslovakia ).

Characters

OTIPs

Crete

Centra] Greece

Yugoslovia

Czechoslovakia

HB

62.0

66.9

67.8

63.5

T

37.3

41.6

39.0

40.2

FA

55.5

57.8

58.2

57.4

E

22.5

24.0

24.7

24.3

GLS

21.6

22.6

22.9

22.4

CL

19.5

20.4

20.2

19.9

ZB

12.1

12.4

12.2

12.2

PC

2.6

2.7

2.8

2.8

CM3

15.3

15.9

16.0

15.7

See Table 1 for key to characters.

Rhinolophus ferrumequinum martinoi Petrov, 1941

Holotype. — Adult female, no. 214, collection of B. Petrov; obtained in April 1939 at Trifunovicevo Brdo, Pepeliste, Yugoslavia.

Distribution. — Southeastern Yugoslavia southward through continental Greece, and on Sicily.

Diagnosis. — Larger than R. f. ferrumequinum. Mean length of forearm varying from 57.71 to 58.16 in samples from Greece; mean condylobasal length in the same samples varying from 20.21 to 20.51; mean zygomatic breadth varying from 12.23 to 12.54.

Remarks. — Compared with R. f. ferrumequinum , the subspecies martinoi is larger in most mensural variates. Of the following measurements (type first, then mean values of topotypes with extremes in parentheses), only five measurements of the holotype are greater than the average measurements of four topotypes (Petrov, 1941): length of head and body, 67.0, 69.37 (68.0-71.5); length of tail, 36.0, 36.25 (35.0-38.0); length of hind foot, 12.0, 12.17 (12.0-12.5); length of ear, 23.7, 23.45 (22.5-24.0); length of forearm, 58.5, 56.27 (54.4-58.2); condylobasal length, 20.8, 20.6 (19.6-21.4); zygomatic breadth, 12.6, 12.57 (12.5-12.7); postorbital constriction, 2.4, 2.45 (2. 2-2. 6); length of maxillary toothrow, 9.0, 9.0 (9. 1-9.6); length of mandibular toothrow, 9.6, 9.6 (9.5-9. 7); length of mandible, 16.2, 15.82 (15.2-16.2).

Specimens examined (total 120, all from Greece). — Thrace: Alexandroupolis, Cave Avantos, 19- Macedonia: Kastoria, Cave Drakos, 3(5, 4$. Thessaly: Elasson, Cave Pythion, 1$. Sterea Hellas: Mesologgi, Vil. Thermos, 2<5; Mesologgi, Cave Varasova, 31(5- 29?; Naupactos, Castle of Naupactos, 1$. Peloponnesus: Achaia, Cave Limnon, 2<5, 1$; Tripolis, Nea Chora, Cave Ag. Heleousa, 14(5, 319-

ILIOPOULOU-GEORGl JDAKI AND ONDRIAS — HORST SHOT BA 1

7

Rhinolophus ferrumequinum ere ti cum , new sul)spec ics

Holotype. — Adult female, skin anti skull no. 5558 ZMI P, obtained on 23 February 1974, in Cave “Milatos,” Lasithi, Crete, by J. Iliopoulou-Georgudaki.

Distribution. — Known only from the Island of Crete.

Diagnosis. — A small (see Table 1), pale-colored representative of R. ferrumequinum. Measurements of the holotype, followed by the mean for 15 individuals from Crete (extremes in parentheses) are: total length, 104.0, 99.30 (92.2-105.0); length of head and body, 63.0, 62.0 (58.4-67.0); length of tail, 41.0, 37.3 (32.6-41.0); length of hind foot, 11.0, 10.61 (9.0-11.4); length of ear, 24.0, 22.46 (20.2-26.0); length of forearm, 58.0, 55.51 (53.3-58.3); greatest length of skull, 21.7, 21.3 (21.2-22.0); condylobasal length, 19.6, 19.55 (18.9-20.4); zygomatic breadth, 12.15, 12.10 (11.6-12.6);

breadth of braincase, 8.5, 8.82 (8.35-9.20); postorbital constriction, 2.4, 2.59 (2.35-2.80); length of maxillary toothrow, 8.2, 7.97 (7.8- 8.0); length of mandibular toothrow, 9.0, 8.98 (8. 7-9.2); length of mandible, 15.4, 15.31 (15.0-16.0).

Remarks. — Compared with R. f. ferrumequinum and R. f. martinoi, the new subspecies creticum is smaller in most measurements and paler in color. The name creticum alludes to the distribution of this new race.

Specimens examined (total 17, all from Crete). — Cave “Milatos,” Lasithi, 1$; Cave “Labyrinth,” Heraclion, 2(5, 19; Starochori Vi 1. , Setia, 5(5, 79. 1 juv.

Summary

In this study, the taxonomic status of specimens of Rhinolophus ferrumequinum from continental Greece and the island of Crete were investigated, using biostatistical methods. The distribution of R. f. martinoi in continental Greece is confirmed, whereas a new subspecies, R. f. creticum, is recognized as occurring on Crete.

Literature Cited

Bate, D. M. A. 1906. On the mammals of Crete. Proc. Zool. Soc. London, pp. 315-523 (for 1905).

Chaworth-Musters, f. 1932. A contribution to our knowledge of the mammals of Macedonia and Thessaly. Ann. Mag. Nat. Hist., set. 10, 9:166-171.

deBeaux, O. 1928. Mammiferi in research faunistich nelle isole Italiane dell’Egeo. Arch. Zool. Italiano, 12' 13:135-153.

Felten, LL, F. Spintzenberger, and G. S torch. 1977. Zur Kleinsaugerfauna West Anatoliens. Senckenberg. Biol. , 58: 1 -44.

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OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY

Festa, E. 1914. Escursioni zoologische de Dr. Enrico Fest nell Isola di Rodi, mammiferi. Boll. Mus. Zool. Anat. Comp., Torino, 29(868): 1-21.

Hanak, V. 1964. Zur Kenntnis der Fledermaus-Fauna Albaniens. Acta Soc. Zool. Bohemoslovenicae, 28:68-88.

Kahmann, H. 1959. Beitrag zur Kenntnis der Fledermaus-Fauna auf der Insel Kreta. Sauget. Mitt., 7:153-156.

Lindberg, K. 1955. Notes sur les grottes de la Grece. Acta Musei Maced. Sci. Nat., 3, 2/14:41-69.

Miller, G. S., Jr. 1912. Catalogue of the mammals of Western Europe. British Museum, London, xv+1019 pp.

Miric, C. 1960. Die Fledermause der unterirdischen Gauge der Festunp Petrovaradin. Bull. Mus. Hist. Nat. Beograd, 16:145-175.

Neithammer, J. 1962. Die Saugetiere von Korfu. Bonner Zool. Beitr., 13:1-49. Petrov, B. 1941. Zamjetki po sistematike i ekologiji melkopitajuscih Juznoj Serbiji. Zap. Russ. Nauk. Inst., Beograd, 16:57-64 (for 1940).

Pieper, H. 1977. Fledermause aus Schleiereulen-Gewollen von der insel Kreta. Z. Saugetierk., 42:7-12.

Pohle, H. 1953. Uber Fledertiere von Kreta. Z. Saugetierk., 17:14-20.

Ridgway, R. 1885. A nomenclature of colors for naturalists. Privately printed, Washington, D.C.

Address of authors: Department of Biology, University of Patras, Patra, Greece. Received 20 March 1985, accepted 2 April 1985.

OCCASIONAL PAPERS

THE MUSEUM

TEXAS TECH UNIVERSITY

NUMBER 103

22 SEPTEMBER 1986

SYNOPSIS OF THE RICE RATS (GENUS ORYZOMYS)

OF NICARAGUA

J. Knox Jones, Jr., and Mark D. Engstrom

Rice rats of the cricetine genus Oryzomys occur from the southeastern United States southward to southern South America; they represent a taxonomically complex group in which a varying number of subgenera are recognized, depending on authority consulted. T he genus evidently first was reported from Nicaragua by Oldfield Thomas (1895:57), who recorded both Oryzomys couesi and O. gracilis ( —O . alfaroi) from Managua. Later, C. Hart Merriam (1901:284) named and described Oryzomys richmondi, now a synonym of O. couesi, and Thomas (1905:586) described the distinctive O. dimidiatus from the Escondido River near Rama. Subsequently (see especially J. A. Allen, 1908, 1910), five additional species of Oryzomys have been found to occur in the country, and we here provide a synopsis of the distribution, systematics, and natural history of those taxa currently known from Nicaragua.

Although there is a diversity of rice rats in Nicaragua (eight species representing at least four subgenera), no species, with the possible exception of O. dimidiatus, is endemic to the country. Nicaraguan Oryzomys comprise a mixture of taxa that are distributed primarily in northern Middle America (O. alfaroi, O. couesi, O. fulvescens, and O. melanotis) with others that occur primarily in southern Central America and adjacent South America (O. alfan, O. bombycinus, O. caliginosus, and possibly O. dimidiatus). Other than dimidiatus, only one species, melanotis, might reach distributional limits (southern) in Nicaragua.