Corresponding author: Vladimir A. Lukhtanov (
Academic editor: N. Shapoval
Lukhtanov VA (2014) Chromosome number evolution in skippers (Lepidoptera, Hesperiidae). Comparative Cytogenetics 8(4): 275–291. doi:
The main karyotypic features of organisms, particularly the number of chromosomes, tend to be stable within species (
In contrast to chromosomal conservatism, chromosomal instability characterizes situations where multiple closely related taxa (populations, subspecies and/or species) belonging to a single phylogenetic lineage differ drastically from each other by major chromosomal rearrangements, sometimes resulting in high variability in chromosome number.
Both phenomena - chromosomal conservatism and chromosomal instability - are clearly expressed in insects of the order
Skippers (the family
Here I provide a first world-wide survey of chromosome numbers in skippers based on intensive search and analysis of published data.
The results of literature search are presented in the Table below. It includes all the discovered chromosome counts except n=13 for
The classification of skippers accepted in this paper follows Warren and colleagues (
The table gives the chromosome numbers of 205 species of skippers, i.e. about 5% of the species of the world fauna. This number is not enough to infer any final statements about peculiarities of chromosome numbers distribution within the
Subfamilies
Within the subfamily
In the family
Within the subfamily
The overall evidence indicates that chromosome numbers of
Several groups of skippers display extreme chromosome number variations at the within-species level (Table). The most extreme variations in number of chromosome elements were observed in first meiotic metaphase of
Second, this kind of variation can be caused by violations in meiotic chromosome pairing resulting in appearance of univalents (instead of bivalents) in meiotic prophase (
Between-species variation exists in numerous genera of skippers (Table 1) and is especially expressed in the Nearctic genus
Chromosome number of skippers (
# | Species | Haploid chromosome number | Country | Reference |
---|---|---|---|---|
|
||||
1 | 29 | Japan |
|
|
31 (2n=62) | Japan |
|
||
2 | 31 | Taiwan |
|
|
3 | 31 | Japan |
|
|
31 | Japan |
|
||
4 | 30 | Oman |
|
|
5 | 31 | Madagascar |
|
|
6 | 23 | Madagascar |
|
|
7 | 31 | Madagascar |
|
|
8 | 31 | Madagascar |
|
|
|
||||
9 | 29 | USA (Texas) |
|
|
10 | 31 | USA (Connecticut) |
|
|
11 | 16 | Mexico |
|
|
12 | 31 | Bolivia |
|
|
13 | 31 | Peru |
|
|
14 | 31 | Brazil |
|
|
15 | 25 | Guatemala |
|
|
16 | 31 | Costa Rica |
|
|
31 | Peru |
|
||
31 | Brazil |
|
||
17 | 20, 21 | Brazil |
|
|
18 | 31 | Mexico |
|
|
31 | Mexico |
|
||
31 | USA (Texas) |
|
||
19 | 22 | Brazil |
|
|
20 | 31 | Argentina |
|
|
21 | 31 | USA (Florida) |
|
|
22 | ca 29–30 | Guatemala |
|
|
23 | 31 | Bolivia |
|
|
24 | 16 | Brazil |
|
|
25 | 15 | Brazil |
|
|
26 | 31 | USA (Connecticut) |
|
|
27 | 29 | Brazil |
|
|
28 | 31 | Mexico |
|
|
29 | 14 | Costa Rica |
|
|
30 | 31 | Bolivia |
|
|
31 | Mexico |
|
||
31 | USA (Florida) |
|
||
31 | 31 | Argentina |
|
|
32 | 31 | Argentina |
|
|
|
||||
|
||||
33 | 40 | Brazil |
|
|
34 | (?) |
32(?) | Brazil |
|
35 | 27 | Brazil |
|
|
36 | 28 | Ecuador |
|
|
28 | Colombia |
|
||
37 | 21 | Colombia |
|
|
38 | 28 | Colombia |
|
|
39 | 28 | Brazil |
|
|
40 | 14(?) | Brazil |
|
|
41 | 28 | Argentina |
|
|
42 | 15 | Brazil |
|
|
43 | 30 | Brazil |
|
|
|
||||
44 | 30 | Japan | ||
45 | 30 | Taiwan |
|
|
46 | 30 | Uganda |
|
|
47 | 30 | Kenya |
|
|
48 | 28 | Kenya |
|
|
|
||||
49 | 29 | Senegal |
|
|
|
||||
50 | 31 | Croatia |
|
|
51 | 43–47 | Spain |
|
|
40–52 | France |
|
||
38–46 | Italy |
|
||
52 | 37–48 (with Us) | Iran |
|
|
53 | 32–41 (with Us) | France (Cauterets) |
|
|
54 | 42–58 (with Us) | Italy |
|
|
55 | 30 | France (Salau, Ariege) |
|
|
56 | 31–32 | Lebanon |
|
|
30 | Turkey (Van) |
|
||
30–37 (with Us) | Turkey (Amasya) |
|
||
57 | 30 | Lebanon |
|
|
30 | Turkey |
|
||
58 | 34 | USA (Texas) |
|
|
59 | 30 | Lebanon |
|
|
60 | 30 | Lebanon |
|
|
61 | 30 | Spain |
|
|
30 | Lebanon |
|
||
62 | 29 | ?USA | Lorković in |
|
63 | 30 | Croatia |
|
|
31 | Turkey |
|
||
64 | 31 | Turkey |
|
|
65 | 31 | Slovenia |
|
|
|
||||
66 | 31 | Mexico |
|
|
32 | USA (Texas) |
|
||
67 | 31 | Trinidad |
|
|
68 | 31 | Argentina |
|
|
69 | 31 | Argentina |
|
|
70 | 31 | USA (Connecticut) |
|
|
71 | 31 | Argentina |
|
|
72 | 31 | USA (Florida) |
|
|
73 | 30 | USA (Connecticut) |
|
|
74 | 30 | USA (Connecticut) |
|
|
75 | 31 | USA (Connecticut) | ||
76 | 31 | Lebanon |
|
|
77 | 31 (2n=62) | Japan |
|
|
31 | Japan |
|
||
78 | E. |
31 | USA (Connecticut) |
|
79 | 31 | Croatia |
|
|
31 | France |
|
||
31 | England |
|
||
80 | 31 | USA (Texas) |
|
|
81 | 32 | Tobago |
|
|
82 | 31 | Mexico | ||
83 | 31 | Bolivia |
|
|
|
||||
84 | 15 | Bolivia |
|
|
15 | Mexico |
|
||
85 | 34 | Brazil |
|
|
|
||||
86 | 31 | Argentina |
|
|
87 | 31 | Mexico |
|
|
88 | 31 | Argentina |
|
|
89 | 14, 13 | USA (Texas) |
|
|
90 | 30 | Bolivia |
|
|
30 | Mexico |
|
||
91 | 29 | USA (Texas) |
|
|
92 | 29 | USA (Texas) |
|
|
93 | 30 | Argentina |
|
|
94 | 30 | Chile |
|
|
95 | 31 | Guatemala |
|
|
96 | ca 18–21 | Turkey | Lukhtanov and Kandul 1995 (in |
|
97 | 30 (2n=60) | USA (Texas) |
|
|
28 | USA (Texas) |
|
||
98 | 24 | Finland |
|
|
24 | Croatia |
|
||
24 | Turkey | Lukhtanov and Kandul 1995 (in |
||
99 | 27 | France |
|
|
100 | 30 | Argentina |
|
|
101 | 30 | Turkey | Lukhtanov and Kandul 1995 (in |
|
102 | 30 | Italy |
|
|
103 | 30 | Italy |
|
|
104 | 29 | Italy |
|
|
105 | 30 | France (Peyreleau, Aveyron) |
|
|
106 | 30 | Chile |
|
|
107 | 31 (2n=62) | Japan |
|
|
108 | 31 | Finland |
|
|
33 | England |
|
||
109 | 30 (2n=60) | USA (Texas) |
|
|
32 | USA (Texas) |
|
||
110 | 30 | France |
|
|
111 | 30 | France |
|
|
112 | 31 | Brazil |
|
|
|
||||
113 | 29 | Chile |
|
|
|
||||
114 | 31 | Australia |
|
|
|
||||
|
||||
115 | 24 | Mexico |
|
|
116 | 38 | USA (Arizona) |
|
|
117 | 5 | Mexico |
|
|
118 | 15 | USA (Arizona) |
|
|
119 | 18 | USA (Texas) |
|
|
120 | 9 | USA (Texas) |
|
|
9 | Mexico |
|
||
121 | 15 | USA (Arizona) |
|
|
122 | 21 | USA (Texas) |
|
|
123 | 22 | USA (Texas) |
|
|
22 | USA (Texas) |
|
||
22 | USA or Mexico |
|
||
22 | USA (Texas) |
|
||
124 | 20 | Mexico |
|
|
125 | 10 | USA (Texas) |
|
|
10 | USA (Texas) |
|
||
126 | 10 | USA (Arizona) |
|
|
127 | 9 | Mexico |
|
|
128 | ca27 | Ecuador |
|
|
129 | 10 | Uganda |
|
|
130 | 28 | Brazil |
|
|
131 | 29 | Malaysia |
|
|
132 | ca28 | Argentina |
|
|
133 | 27 | USA |
|
|
134 | 27 | USA (South central Texas) |
|
|
27 | USA (Western Texas) |
|
||
27 | USA |
|
||
27 | USA (Texas) |
|
||
27 | USA (Texas) |
|
||
27 | USA |
|
||
27 | ?Mexico |
|
||
135 | 27 | USA | ||
136 | 26 | USA (Florida) |
|
|
137 | 17 | Uganda |
|
|
138 | 50 | USA (Texas) | ||
139 | 23 | Taiwan |
|
|
140 | 31 (2n=62) | Japan |
|
|
141 | 31 | Japan |
|
|
|
||||
142 | 41 | Yemen |
|
|
41 | Turkey |
|
||
41 | Lebanon |
|
||
143 | 15 | Egypt |
|
|
15 | Israel | |||
144 | 24 | France |
|
|
24 | Alger |
|
||
145 | 16 | Japan | ||
16 | China |
|
||
146 | 16 | Hong Kong |
|
|
147 | 16 (2n=32) | Japan |
|
|
148 | 16 | Japan |
|
|
149 | 16 | Lebanon |
|
|
150 | 16 | Taiwan |
|
|
151 | 16, 17, 18 (2n=32, 33) | Japan |
|
|
16 | Japan |
|
||
152 | 16 | Uganda |
|
|
|
||||
153 | 28 | Australia |
|
|
154 | 29 (2n=58) | Japan |
|
|
155 | 29 | Taiwan |
|
|
156 | 29 | Japan (Okinava) |
|
|
157 | 29 (2n=58) | Taiwan |
|
|
|
||||
158 | 29 | USA (Florida) |
|
|
159 | 27 | England |
|
|
160 | 10 (2n=20) | Japan |
|
|
161 | 28 | Spain |
|
|
162 | 29 | Lebanon |
|
|
163 | 9 (2n=18) | Japan |
|
|
164 | 29 | Finland |
|
|
29 | Lebanon |
|
||
|
||||
165 | 29 | Mexico |
|
|
166 | 30 | Brazil |
|
|
167 | 29 | USA (Texas) |
|
|
168 | 29 | USA (Texas) |
|
|
169 | 29 | USA (Florida) |
|
|
170 | 29 | USA (Texas) |
|
|
|
||||
171 | 28, 29 | USA (Texas) |
|
|
172 | 29 | USA (Texas) |
|
|
173 | 29 | USA (Texas) |
|
|
174 | 29 | USA (Texas) |
|
|
175 | 29 | USA (Texas) |
|
|
176 | 29 | USA (Connecticut) |
|
|
177 | 31 | Tobago |
|
|
178 | 29 | Ecuador |
|
|
179 | 29 (2n=58) | USA (Texas) |
|
|
29 | USA (Texas) |
|
||
180 | 27 | Peru |
|
|
181 | 30 | USA (Connecticut) |
|
|
182 | 25 | Brazil |
|
|
183 | 31, ca32 | Ecuador |
|
|
184 | 26 | Brazil |
|
|
185 | 26 | Brazil |
|
|
|
||||
186 | 48 | USA (Florida) |
|
|
187 | 29 | Argentina |
|
|
188 | 28 | Italy |
|
|
28 | Lebanon |
|
||
189 | 28 (2n=56) | Japan |
|
|
190 | 29 | Argentina |
|
|
191 | 29 | Argentia |
|
|
29 | USA (Florida) |
|
||
192 | 29 | Chile |
|
|
193 | 29 (2n=58) | Japan |
|
|
24 | Japan |
|
||
194 | 29 | USA |
|
|
195 | 29 | Finland |
|
|
29 | Croatia |
|
||
196 | 29 (2n=58) | Japan |
|
|
197 | 29 | USA (Florida) |
|
|
198 | 29 | ?USA | Lorković in |
|
199 | 29 | USA |
|
|
200 | 29 | USA (Connecticut) |
|
|
201 | 29 | USA (Florida) |
|
|
202 | 29 | Argentina |
|
|
27 | USA (Texas) |
|
||
29 | USA (Florida) |
|
||
203 | 28 | USA (Texas) |
|
|
204 | 28–30 | USA (Texas) |
|
|
205 | 27 | Argentina |
|
Recent years karyological data have been widely used in studies of butterfly taxonomy and in biodiversity research as main or additional chracters for detecting cryptic species (e.g.
In the genus
Thus, interspecific chromosomal differences are useful for discovering and describing new cryptic species of
I thank A. Warren (University of Florida) for help and consultations in taxonomy and nomenclature of skippers. The study was supported by the Russian Foundation for Basic Research: mainly by grant RFBR 13-04-92716-IND-a and partially by grant RFBR 14-04-01051-a.