Alfonso N. García Aldrete and J. Arturo Casasola Gonzalez
Instituto de Biología
UNAM. Departamento de Zoología
Apartado Postal 70- 153
04510 Mexico, D.F. Mexico

Abstract

A survey of the Psocoptera of the Calakmul Biosphere Reserve, Campeche, Mexico, was conducted in 1997 and early 1998. The collecting effort was 260 man-hours, excluding the operation of light and Malaise traps. A total of 1675 specimens was taken, representing 96 species, in 48 genera and 23 families. The µ Diversity Index for this collection was 22.12. Fifteen species constituted 66.7% of the total number of specimens, and 40 species constituted 3.9% of the same total. Only 18 of the 96 species present in the area are widely distributed locally, whereas 72 of the 96 species in the area showed restricted local distribution. The level of endemism is high (19.79% of the total number of species).

Key Words: Calakmul Biosphere Reserve, Campeche, Mexico, Psocoptera

Resumen

Durante 1997 y principios de 1998 se condujo un censo de Psocoptera en la Reserva de la Biósfera de Calakmul, Campeche, en el que el esfuerzo de colecta fue de 260 horas-hombre, sin contar el tiempo de operación de trampas de luz y trampas Malaise. Fueron capturados un total de 1675 ejemplares, que representan a 96 especies, en 48 géneros y 23 familias. El Indice de Diversidad µ, calculado para ésta colección, fue de 22.12. Quince especies constituyeron el 66.7% del total de ejemplares recolectados, mientras que 40 especies constituyeron 3.9% del mismo total. Sólo 18 de las 96 especies registradas en la área tienen una amplia distribución local, y 72 del total de 96 especies tienen una distribución local muy restringida. El nivel de endemismo es alto (19.76% del total de especies).

The Calakmul Biosphere Reserve, in the Mexican state of Campeche, was created on 22 May, 1989, by decree of the then President of Mexico, Carlos Salinas de Gortari. The reserve is located at the base of the Yucatan Peninsula, in the southwestern corner of Campeche, between 17°49' and 19°11'N and between 89°08' and 90°08'W, bordering on the south with the Guatemalan Peten and partially to the east with the state of Quintana Roo. It covers approximately 7000 square kilometers, or about 14% of the total area of Campeche. It has a peculiar shape (Fig. 1), with two large areas, one to the north and one to the south of the highway Escarcega-Chetumal, separated by a pronounced narrowing that crosses the highway some 15 kilometers west of X'puhil. The defects in the design of the reserve have been widely pointed out and discussed by Galindo Leal (1997). All in all, it constitutes the largest humid forest reserve area in the country, with representation, in order of importance of area covered, of medium subperennifolious forest, low subperennifolious forest, secondary vegetation, perennifolius-subperennifolious evergreen forest, and aquatic vegetation (Gomez Pompa & Dirzo 1995). The area is inhabited by many species of wild animals, threatened or in danger of extinction, such as jaguar, ocelot, jaguarundi, spider and howler monkeys, curassow, harpy eagle and tapir. The area is also rich in Mayan archaeological zones of the Classic period, in architectural styles Peten, Chenes and Rio Bec (e.g. Calakmul, Hormiguero, Chicanna, Becan and Balamkum).

With respect to the Psocoptera fauna, the only notable reference is the record, by Mockford & Garcia Aldrete (1996), of 26 species in Campeche, which were the result of isolated, not systematic insect collecting in several localities in the state, none of these in the reserve area, with only some records from the vicinities of X'puhil. Most of the species recorded were neotropical or pantropical, widely distributed and also occurring in the Caribbean.

The purpose of this work was to survey the fauna of psocids in the reserve area and surroundings, to estimate the relative abundance and local distribution of the species present, and to determine the specific richness of the different sites sampled. The specimens collected are deposited in the National Collection of Insects (Departamento de Zoologia, Instituto de Biologia, UNAM, Apartado Postal 70-153, 04510, Mexico, D.F.)

Materials and Methods

In May and September, 1997, and in February, 1998, psocid collecting was conducted in the reserve area and some neighbouring places. The insects were taken by beating the vegetation, sifting litter, directly examining tree trunks and rock faces, and by using light and Malaise traps. During the first collecting event (1-9.V.1997), the effort was of 135 man-hours, then 70 man-hours during the second collecting event (19-25.IX.1997), and 55 man-hours during the third collecting event (15-19.II.1998). The specimens collected were preserved directly in 80% ethanol. Table 1 presents a list of the collecting localities and their geographic coordinates, and they are also indicated in Figure 1. It is pertinent to point out that no collecting was done in the northern segment of the reserve, nor in the nuclear zones.

Results

During the first collecting event, 708 psocid specimens were taken, with 58 species being represented. During the second collecting event, 449 specimens were taken, representing 41 species, 16 of which had not been taken during the first event, and during the third collecting event, 518 specimens were taken, representing 66 species, 22 of which had not been previously collected. A total of 1675 specimens was collected, representing 96 psocid species, in 48 genera and 23 families (Table 2).

Figure 2 shows the species accumulation curve for the collecting period. The slope of the line indicates that a fourth collecting episode would have been needed to determine if the curve was or was not in the asymptotic phase. With the evidence that in the third collecting event 22.9% of the total number of species were new additions, it is likely that more unrecorded psocid species could still be found in the area.

Table 1. Collecting localities in the Calakmul Biosphere Reserve and vicinity.

1.
25 km N of Calakmul archaeological zone, 230 m.
18°17'49"N, 89°50'36"W
2.
Calakmul archaeological zone, ca. large "aguada", 265 m.
18°07'26"N, 89°48'56"W
3.
Calakmul archaeological zone, 265 m.
18°06'35"N, 89°48'17"W
4.
El Chorro, ejido Nuevo Becal, 130 m.
18°35'25"N, 89°15'28"W
5.
Laguna de Alvarado, 316 m.
18°01'54"N, 89°15'45"W
6.
Laguna de Alvarado, 322 m.
18°00'55"N, 89°16'10"W
7.
Hormiguero archaeological zone, 295m.
18°24'10"N, 89°29'13"W
8.
Arroyo Colon, ejido C. Colon, 420 m.
18°12'59"N, 89°27'23"W
9.
San Antonio Soda, ejido Diaz Ordaz, 200 m.
18°24'54"N, 89°08'19"W
10.
Zoh Laguna, ca. "aguada", 327 m.
18°35'21"N, 89°25'07"W
11.
La Mancolona, ejido 20 de Junio, 232 m.
18°48'38"N, 89°17'29"W

Table 2 lists the species of psocids collected in the area, the species and number of specimens taken in each collecting event, the relative abundance of each species, the amplitude of distribution in the area sampled (A = number of localities in which each species was found), and the hierarchic order of each species (HOS), an ordering in which the species are placed in hierarchy, according to their importance values; in this case, the number of specimens/species was taken as importance value.

The 96 species found represent 48 genera in 23 families. The genus most diverse is Lachesilla, with 13 species, followed by Tapinella, Caecilius and Archipsocus, each with five species; then follow Echmepteryx, Lithoseopsis and Peripsocus, with four species each, and Psyllipsocus, Liposcelis, Ectopsocus, Archipsocopsis, Blastopsocus and Ptycta, with three species each. The genera Cladiopsocus, Hemipsocus and Trichadenotecnum are represented by two species each, and there is a large group of 32 genera represented by only one species each.

In terms of relative abundance, the 96 species are distributed in 38 ranks of hierarchic importance (Fig. 3). The species numerically most important is Archipsocopsis sp. 1, with 209 specimens, followed by Ectopsocus titschacki Jentsch, with 108 specimens, Echmepteryx alpha Garcia Aldrete, with 92 specimens, Hemipsocus africanus Enderlein, with 86 specimens, and Caecilius totonacus Mockford, with 78 specimens. Together, the 15 most abundant species constitute 66.7% of the total number of individuals, and, on the opposite end, 19 species are represented by one specimen, 16 species are represented by two specimens, and five species are represented by three specimens, so that 40 species constitute only 3.9% of the total of specimens collected.

The µ Diversity Index [S = µ log (1 + N/µ), cf. Taylor, Kempton & Woiwod (1976)], calculated for the Calakmul psocid collection, resulted in a value of 22.12, one of the highest recorded in the literature, surpassed only by the diversity indices for the Psocoptera of Chamela, Jalisco, Mexico (µ = 24.01, N = 2863, S = 115), Panama Lowlands (µ = 24.5, N = 10092, S = 148), and Los Tuxtlas, Veracruz, Mexico (µ = 32.45, N = 4194, S = 158) (Broadhead & Wolda 1985; Garcia Aldrete 1988; Garcia Aldrete, Mockford & Garcia Figueroa 1997).

Table 3 presents the species and number of specimens of each species collected in each locality during this study; it also includes the habitats in which the species were collected. Since the collecting effort was not the same in each locality, the results are biased; however, the comparatively high species richness of localities 3, 7 and 10 probably reflect also intrinsic differences among the localities sampled. The richer ones are sites physically complex, varied, with several habitats sampled, such as the Calakmul archaeological zone, the Hormiguero archaeological zone or Laguna de Alvarado.

The species of psocids collected in the Calakmul area, can be assigned to the following biogeographic categories:

I. Endemics and presumed endemics (19 species).

Nepticulomima sp., Rhyopsocus sp., Psyllipsocus sp. 2, Lithoseopsis sp. 4, Liposcelis sp., Tapinella sp. 1, Xanthocaecilius sp., Nanolachesilla sp., Peripsocus sp. 4, Archipsocopsis sp. 3, Palmicola sp., Blastopsocus spp.1, 2, and 3, Metylophorus sp., Steleops sp., Ptycta sp.1, and Trichadenotecnum spp. 1 and 2.

II. Tropical waifs (9 species).

Proentomum personatum Badonnel, Soa flaviterminata Enderlein, Echmepteryx falco Badonnel, E. madagascariensis (Kolbe), Nanopsocus oceanicus Pearman, Ectopsocus titschacki Jentsch, E. vilhenai Badonnel, Pseudocaecilius citricola (Ashmead) and Hemipsocus africanus Enderlein.

III. Cosmopolitan species (2 species).

Liposcelis bostrychophila Badonnel, Caecilius casarum Badonnel.

IV. Species widespread in tropical and subtropical America (9 species).

Thylacella cubana (Banks), Belaphopsocus badonneli New, Liposcelis ornata Mockford, Tapinella maculata Mockford & Gurney, Dasypsocus roesleri (New & Thornton), Lachesilla cuala Garcia Aldrete, Peripsocus potosi Mockford, Nepiomorpha brasiliana Badonnel, and Cerastipsocus trifasciatus (Provancher).

V. Species occurring in Mexico and southeastern USA (2 species).

Lachesilla bottimeri Mockford & Gurney, L. penta Sommerman.

VI. Species occurring in tropical Mexico and Guatemala or Belize, not extending to Central America and the Caribbean (7 species).

Echmepteryx alpha Garcia Aldrete, Cladiopsocus garciai Eertmoed, Triplocania spinosa Mockford, Anomopsocus sp. a, Lachesilla disjuncta Garcia Aldrete, L. nuptialis Badonnel & Garcia Aldrete, Ptycta tikala Mockford.

VII. Species occurring in tropical Mexico, Central America and the Caribbean (2 species).

Lachesilla denticulata Garcia Aldrete, L. riegeli Sommerman.

VIII. Species occurring in tropical Mexico and the Caribbean (5 species).

Echmepteryx intermedia Mockford, Neolepolepis caribensis (Turner), Tapinella olmeca Mockford, Lachesilla yanomamioides Garcia Aldrete, Hemipsocus pretiosus Banks.

IX. Species occurring in tropical Mexico and Central America (5 species).

Caecilius totonacus Mockford, C. sp. 3, Lachesilla tropica Garcia Aldrete, Scytopsocus ca. coriaceous Roesler, Lichenomima varia (Navas).

X. Species restricted to the Yucatan Peninsula (5 species).

Psyllipsocus spp. 1 and 3, Lithoseopsis sp. 1, Loneura leonilae Garcia Aldrete, Ptycta sp. 2.

XI. Species occurring in the Yucatan Peninsula and neighbouring areas (6 species).

Lithoseopsis spp. 2 and 3, Archipsocus sp. 1, Heterocaecilius badonneli Garcia Aldrete, Aaroniella sp., Myopsocus sp.

XII. Species occurring in tropical Mexico (20 species).

Electrentomopsis variegatus Mockford, Tapinella vittata Garcia Aldrete, Pachytroctes ixtapaensis Garcia Aldrete, Epipsocus sp., Dolabellopsocus roseus Eertmoed, Cladiopsocus ocotensis Garcia Aldrete, Caecilius sp. 2, Lachesilla bifurcata Garcia Aldrete, L. sp. (forcepeta group), L. pedicularia group, Ectopsocus mexicanus Garcia Aldrete, Peripsocus chamelanus Badonnel, P. ca. stagnivagus Chapman, Archipsocopsis spp. 1 and 2, Archipsocus spp. 2, 3, 4, and 5, Haplophallus sp.

XIII. Species occurring in Cuba (2 species).

Embidopsocus cubanus Mockford, Pseudarchipsocus guajiro Mockford.

XIV. Species restricted to Guatemala or Belize (3 species).

Tapinella sp. 2, Notiopsocus sp., Lachesilla F9B.

Given the geographic location of Calakmul, the composition of its psocid fauna does not contain elements of surprise and it is rather as expected for an area near the edge of tropical Mexico, and close to Central America and the Caribbean; it is dominated by Mexican tropical species, with the addition of the species widespread in tropical America, plus the species also shared with Central America and the Caribbean region, plus the usual array of tropical waifs and cosmopolitans. Categories IX and X, of species restricted to the Yucatan Peninsula or occurring nearby, point to the biotic distinctness of that area (see also Barrera 1962). The category of endemics, comprising 19.79% of the fauna of Calakmul, gives it the element of uniqueness. It is pertinent to note that 18 of the 26 species previously recorded in Campeche, were found in the area of the Calakmul Reserve.

The results of this survey indicate that the psocid community of the Calakmul Biosphere Reserve area is rich in species, with a high proportion of endemics. It also indicates that the community shows fragility in that there is a large number of "rare" species (e.g. 40 species of which only 1-3 specimens were collected throughout the sampling period), and in that a large number of species have only a small amplitude of local distribution (e.g. 72 species collected in only one or two localities), with which environmental changes, either natural or anthropogenic, could result in local extinctions.

Acknowledgements

This project is part of a larger one, financed by the Mexican agency CONABIO (Project M003 "Reconocimiento de la biodiversidad de la Reserva de la Biósfera Calakmul: Odonata, Psocoptera y Diptera Acuáticos (Insecta)"). Atilano Contreras, Enrique Gonzalez, Tomas Martinez, Adolfo Ibarra, and Rocio Lopez participated in it and contributed with specimens of Psocoptera. To all of them, and to CONABIO, our most sincere thanks.

References Cited

Barrera, A. 1962. La península de Yucatán como provincia biótica. Revista de la Sociedad Mexicana de Historia Natural 23: 71-105.

Broadhead, E., and H. Wolda. 1985. The diversity of Psocoptera in two tropical forests in Panama. Journal of Animal Ecology 54: 739-754.

Galindo-Leal, C. 1997. Diseño de reservas: el "mal congénito" de Calakmul. Ecotono. Centro para la biología de la conservación. Boletín del Programa de Investigación Tropical. Stanford University. Pp.4-7.

Garcia Aldrete, A. N. 1988. The psocids (Psocoptera) of Chamela, Jalisco, Mexico. Species, diversity, abundance distribution and seasonal changes. Folia Entomologica Mexicana 77: 63-84.

Garcia Aldrete, A. N., E. L. Mockford, and J. Garcia Figueroa. 1997. Psocoptera, p. 299-309 in Gonzalez Soriano, E., R. Dirzo, and R. Vogt. Historia Natural de Los Tuxtlas. Instituto de Biología-Instituto de Ecología, UNAM. Mexico.

Gomez Pompa, A., and R. Dirzo. 1995. Reservas de la biósfera y otras areas naturales protegidas de México. Instituto Nacional de Ecología (SEMARNAP)- CONABIO. Mexico, D. F. 159 pp.

Mockford, E. L., and A. N. Garcia Aldrete. 1996. Psocoptera, p. 175-205 in Llorente, B. J., A. N. Garcia Aldrete, and E. Gonzalez S. Biodiversidad, taxonomía y biogeografía de artrópodos de México. Instituto de Biología, UNAM., Mexico.

Taylor, L. R., R. A. Kempton, and I. P. Woiwod. 1976. Diversity statistics and the log-series model. Journal of Animal Ecology 45: 255-272.

Fig. 1. Location of the Calakmul Biosphere Reserve, Campeche, and Psocoptera collecting localities in the area.

Fig. 2. Species accumulation curve for the Psocoptera of the Calakmul area. May 1997-February 1998.

Fig. 3. Species abundance distribution of the collection of Psocoptera from the Calakmul Biosphere Reserve and surrounding areas. Log. of abundance ranked against species. µ = 22.12.

Table 2. (Continued) Psocoptera from the Calakmul Biosphere Reserve, Campeche and vicinity (N = number of specimens, %T = percentage of the total, A = number of localities in which each species was collected, HOS = hierarchic order of species).

1-9. V. 1997
(135 man hours)
19-25. IX. 1997
(70 man hours)
15-19. II. 1998
(55 man hours)
N
%T
A
HOS
males
females
nymphs
males
females
nymphs
males
females
nymphs
TROGIOMORPHA
Lepidopsocidae
1
Thylacella cubana (Banks), 1941
2
2
1
5
0,30
4
34
2
Nepticulomima Enderlein, 1906
7
14
21
1,25
2
19
3
Proentomum personatum
Badonnel, 1949
7
12
5
10
3
6
43
2,57
10
12
4
Soa flaviterminata
Enderlein, 1906
1
7
1
9
0,54
3
30
5
Echmepteryx alpha Garcia Aldrete, 1984
38
30
1
4
13
6
92
5,49
7
3
6
E. falco Badonnel,1949
4
8
12
0,72
2
27
7
E. madagascariensis (Kolbe), 1885
22
9
3
3
5
24
66
3,94
3
7
8
E. intermedia Mockford, 1974
5
4
1
2
2
3
17
1,01
4
23
9
Neolepolepis caribensis (Turner), 1975
3
3
0,18
1
36
Psoquillidae
10
Rhyopsocus sp.
1
1
0,06
1
38
Psyllipsocidae
11
Psyllipsocus Selys-Longchamps, 1872. sp. 1
12
14
1
5
3
1
36
2,15
3
13
12
P. sp. 2
8
1
4
5
18
1,07
3
22
13
P. sp. 3
1
1
0,06
1
38
TROCTOMORPHA
Amphientomidae
14
Lithoseopsis Mockford, 1993. sp. 1
3
9
2
1
1
4
20
1,19
5
20
15
L. sp. 2
1
1
2
0,12
1
37
16
L. sp. 3
8
4
12
0,72
1
27
17
L. sp. 4
1
1
0,06
1
38
Compsocidae
18
Electrentomopsis variegatus
Mockford, 1967
1
1
2
0,12
1
37
Liposcelididae
19
Belaphopsocus badonneli New, 1971
2
2
0,12
1
37
20
Embidopsocus cubanus Mockford, 1987
1
1
2
0,12
1
37
21
Liposcelis bostrychopila Badonnel, 1931
2
2
0,12
2
37
22
L. ornata Mockford, 1978
2
1
3
0,18
2
36
23
Liposcelis Motschulsky, 1852
1
1
0,06
1
38
24
Nanopsocus oceanicus Pearman, 1928
13
2
6
21
1,25
3
19
25
Tapinella maculata Mockford & Gurney, 1926
2
6
2
2
3
15
0,90
7
24
26
T. olmeca Mockford, 1975
4
25
5
1
6
1
2
44
2,63
6
10
27
T. vittata Garcia Aldrete, 1993
2
17
3
4
2
23
2
53
3,16
6
10
28
Tapinella Enderlein, 1908. sp. 1
15
4
19
1,13
6
21
29
T. sp. 2
8
3
11
0,66
3
28
30
Pachytroctes ixtapaensis Garcia
Aldrete, 1986
1
3
4
0,24
1
35
PSOCOMORPHA
Epipsocidae
31
Epipsocus Hagen, 1866
2
3
1
6
0,36
2
33
Dolabellopsocidae
32
Dolabellopsocus roseus Eertmoed, 1973
1
1
0,06
1
38
Cladiopsocidae
33
Cladiopsocus garciai Eertmoed, 1986
1
4
3
8
0,48
2
31
34
C. ocotensis Garcia Aldrete, 1996
1
4
3
1
1
3
13
0,78
2
26
Ptiloneuridae
35
Loneura leonilae Garcia Aldrete, 1995
1
1
2
0,12
1
37
36
Triplocania spinosa Mockford, 1957
3
5
7
15
0,90
2
24
Asiopsocidae
37
Notiopsocus Banks, 1913
1
3
7
11
0,66
2
28
Caeciliidae
38
Caecilius casarum Badonnel, 1931
2
2
0,12
1
37
39
C. totonacus Mockford, 1966
1
51
27
79
4,72
3
5
40
Caecilius Curtis, 1837. Sp. 1
1
4
4
9
0,54
3
30
41
C. sp. 2
1
1
0,06
1
38
42
Xanthocaecilius Mockford, 1989
1
1
0,06
1
38
Amphipsocidae
43
Dasypsocus roesleri (New & Thornton), 1975
1
1
7
3
13
25
1,49
4
18
Lachesillidae
44
Anomopsocus Roesler, 1940
1
1
0,06
1
38
45
Nanolachesilla Mockford &
Sullivan, 1986
1
1
0,06
1
38
46
Lachesilla bottimeri Mockford & Gurney, 1956
1
1
0,06
1
38
47
L. bifurcata Garcia Aldrete, 1986
1
1
0,06
1
38
48
L. sp. (forcepeta group)
2
5
3
2
3
1
3
13
32
1,91
6
14
49
L. cuala Garcia Aldrete, 1988
2
2
0,12
1
37
50
L. denticulata Garcia Aldrete, 1988
3
2
5
18
35
63
3,76
5
8
51
L. disjuncta Garcia Aldrete, 1988
1
7
16
1
1
1
27
1,61
4
16
52
L. nuptialis Badonnel & Garcia
Aldrete, 1980
5
1
7
5
10
28
1,67
8
15
53
L. penta Sommerman, 1946
1
3
17
2
8
1
3
9
44
2,63
7
11
54
L. riegeli Sommerman, 1946
1
1
1
3
0,18
2
36
55
L. tropica Garcia Aldrete, 1982
3
1
3
1
8
0,48
6
31
56
L. yanomamioides Garcia Aldrete, 1996
2
2
9
4
7
7
6
15
26
78
4,66
6
6
57
Lachesilla Westwood, 1840. sp. F9 B
2
2
7
11
0,66
1
28
58
L. sp. (pedicularia group)
1
2
3
1
4
11
0,66
7
28
Ectopsocidae
59
Ectopsocus mexicanus Garcia
Aldrete, 1991
1
1
0,06
1
38
60
E. titschacki Jentsch, 1929
17
45
12
10
16
2
2
3
1
108
6,45
7
2
61
E. vilhenai Badonnel, 1955
4
6
10
0,60
3
29
Peripsocidae
62
Peripsocus potosi Mockford, 1971
1
3
1
5
0,30
3
34
63
P. chamelanus Badonnel, 1986
1
1
2
0,12
2
37
64
P. ca. stagnivagus Chapman, 1930
1
1
2
0,12
2
37
65
P. sp. 1
1
2
3
0,18
2
36
Archipsocidae
66
Archipsocopsis Badonnel, 1966. sp. 1
1
54
2
8
101
42
1
209
12,48
7
1
67
A. sp. 2
5
1
1
7
0,42
2
32
68
A. sp. 3
1
1
2
0,12
1
37
69
Archipsocus Hagen, 1882 sp. 1
2
2
0,12
1
37
70
A. sp. 2
7
1
4
1
33
11
57
3,40
9
9
71
A. sp. 3
1
1
2
0,12
1
37
72
A. sp. 4
2
2
0,12
2
37
73
A. sp. 5
1
1
0,06
1
38
74
Pseudarchipsocus guajiro
Mockford, 1974
1
2
3
0,18
1
36
Pseudocaeciliidae
75
Pseudocaecilius citricola
(Ashmead), 1879
1
1
3
2
7
0,42
4
32
76
Heterocaecilius badonneli Garcia Aldrete, 1989
4
6
4
14
0,84
1
25
77
Scytopsocus Roesler, 1940 (ca. coriaceous Roesler, 1940)
2
1
1
3
7
0,42
3
32
Philotarsidae
78
Haplophallus Thornton, 1959
2
1
3
6
0,36
3
33
79
Aaroniella Mockford, 1951
1
1
2
0,12
1
37
Elipsocidae
80
Palmicola Mockford, 1955
1
1
2
0,12
2
37
81
Nepiomorpha brasiliana Badonnel, 1973
1
14
15
0,90
2
24
Hemipsocidae
82
Hemipsocus africanus Enderlein, 1907
6
13
7
12
28
14
2
4
86
5,13
5
4
83
H. pretiosus Banks, 1930
2
5
1
8
0,48
1
31
Psocidae
84
Blastopsocus Roesler, 1943. Sp.1
1
2
2
5
0,30
3
34
85
B. sp. 2
1
1
16
18
1,07
2
22
86
B. sp. 3
1
1
0,06
1
38
87
Cerastipsocus trifasciatus
(Provancher), 1876
1
1
1
14
17
1,01
3
23
88
Metylophorus Pearman, 1932
1
1
0,06
1
38
89
Steleops Enderlein, 1910
1
1
0,06
1
38
90
Ptycta Enderlein, 1925. sp. 1
1
1
0,06
1
38
91
P. sp. 2
1
1
0,06
1
38
92
P. tikala (Mockford), 1957
1
1
0,06
1
38
93
Trichadenotecnum Enderlein, 1909. sp. 1
1
1
1
2
5
0,30
2
34
94
T. sp. 2
1
1
2
4
0,24
2
35
Myopsocidae
95
Lichenomima varia (Navas), 1927
15
9
1
1
26
1,55
2
17
96
Myopsocus Hagen, 1866
5
1
3
9
0,54
4
30
TOTAL
76
423
212
68
256
122
50
240
228
TOTAL
711
446
518
1675
a DIVERSITY INDEX (S = a log (1+N/a)) =
22,12

Table 3. (Continued) Psocoptera of the Calakmul Biosphere Reserve, Campeche, and vicinity. number of species taken in each locality, and habitats in which each species was collected. I. Branches and foliage of shrubs. II. Leaf litter. III. Tree trunks and bark. IV. Typha foliage. V. Dead palm fronds. VI. Bromeliads, orchids and other epiphytes. VII. Herbaceous plants. VIII. Calcareous rock faces. IX. Abandoned termite nest. X. Malaise trap. XI. Light trap.

Localities
Habitats
1
2
3
4
5
6
7
8
9
10
11
I
II
III
IV
V
VI
VII
VIII
IX
X
XI
TROGIOMORPHA
Lepidopsocidae
1
Thylacella cubana (Banks), 1941
2
1
1
*
*
*
2
Nepticulomima Enderlein, 1906
17
4
*
*
*
*
3

Proentomum
personatum
Badonnel, 1949
2
11
1
1
12
2
1
1
5
7
*
*
*
*
*
*
*
4
Soa flaviterminata Enderlein, 1906
5
1
3
*
*
5
Echmepteryx alpha Garcia Aldrete, 1984
12
23
10
5
29
6
7
*
*
*
*
*
*
*
*
*
*
*
6
E. falco
Badonnel, 1949
6
6
*
*
*
*
*
7
E. madagascariensis (Kolbe), 1885
14
45
7
*
8
E. intermedia
Mockford, 1974
3
5
5
4
*
*
9
Neolepolepis caribensis (Turner), 1975
3
*
Psoquillidae
10
Rhyopsocus sp.
1
*
Psyllipsocidae
11

Psyllipsocus
Selys-Longchamps, 1872. sp. 1
12
11
13
*
*
12
P. sp. 2
7
9
2
*
13
P. sp. 3
1
*
TROCTOMORPHA
Amphientomidae
14
Lithoseopsis
Mockford, 1993. sp. 1
2
1
13
2
2
*
*
*
15
L. sp. 2
2
*
16
L. sp. 3
12
*
17
L. sp. 4
1
*
Compsocidae
18

Electrentomopsis variegatus Mockford, 1967
2
*
Liposcelididae
19
Belaphopsocus badonneli New, 1971
2
*
20
Embidopsocus cubanus Mockford, 1987
2
*
21
Liposcelis bostrychopila Badonnel, 1931
1
1
*
22
L. ornata Mockford, 1978
2
1
*
*
23
Liposcelis Motschulsky, 1852
1
*
24
Nanopsocus oceanicus Pearman, 1928
18
1
2
*
*
25
Tapinella maculata Mockford & Gurney, 1926
1
5
1
5
1
1
1
*
*
*
26
T. olmeca Mockford, 1975

16
18
2


6

1
1


*



*

*




27
T. vittata Garcia
Aldrete, 1993
4
27
1
8
9
4
*
*
*
*
*
*
28
Tapinella Enderlein, 1908. sp. 1
2
1
1
2
10
3
*
*
29
T. sp. 2
7
1
3
*
*
*
30

Pachytroctes ixtapaensis Garcia Aldrete, 1986
4
*
PSOCOMORPHA
Epipsocidae
31
Epipsocus Hagen, 1866
1
5
*
*
Dolabellopsocidae
32
Dolabellopsocus roseus Eertmoed, 1973
1
*
Cladiopsocidae
33
Cladiopsocus garciai Eertmoed, 1986
7
1
*
*
*
*
34
C. ocotensis Garcia Aldrete, 1996
5
8
*
*
Ptiloneuridae
35
Loneura leonilae
Garcia Aldrete, 1995
2
*
*
36
Triplocania spinosa Mockford, 1957
14
1
*
*
Asiopsocidae
37
Notiopsocus Banks, 1913
9
2
*
*
*
Caeciliidae
38
Caecilius casarum
Badonnel, 1931
2
*
39
C. totonacus
Mockford, 1966
1
69
9
*
*
40
Caecilius Curtis, 1837. sp. 1
6<